Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

A Preliminary World-wide Key to the (including Lepra species)

A.W. Archer & J.A. Elix

The lichen genus Pertusaria () is widely distributed throughout the world, from equatorial to polar regions (Dibben 1980; Lumbsch & Nash 2001). Species may grow on bark, rock, soil, plant débris and mosses and are differentiated by the apothecial structure (disciform or verruciform), the number and structure of the ascospores (1, 2, 4 or 8 per ascus, smooth- or rough-walled ascospores) and the chemistry (Dibben 1980; Archer 1997). Chemistry has been recognised as an important taxonomic tool in the identification of species in the genus Pertusaria (Lumbsch 1998).

The chemistry of the genus Pertusaria has been reported in many publications. Oshio (1968) reported the colour reactions of Japanese species and the compounds producing these colours were subsequently identified by Dibben (1975) who later published the chemistry of North American Pertusaria (1980). Similarly, Poelt and Vĕzda (1981) described the colour reactions of European species of Pertusaria and the identity of these compounds was later determined by Hanko (1983). Additional synonymy and chemical data for a range of European taxa was reported by Niebel-Lohmann and Feuerer (1992). Chemical data on many type specimens was reported by Archer (1993, 1995) and the chemistry of Australian Pertusaria published (Archer 1997). Additional type specimens hace since been examined and their chemistries determined. A current Key to European Pertusaria (Sipman, www.bgbm.org/BGBM/Staff/Wiss/Sipman/keys/perteuro.htm) contains much chemical information on European taxa, which has been included in this Key. Modern descriptions of new taxa in the genus would now be regarded as incomplete without a report of the chemistry.

The total number of species in the genus is not accurately known. Nash et al. (2001) suggest ca. 350 species and American Record Express (http://americanrecordexpress.com/fungi/p/) suggests ca. 920 species. Archer & Messuti (2009) calculated a possible 1550 species out of a theoretically possible 3500 chemical and morphological combinations (Archer 1997) while the Index Fungorum (www.indexfungorum.org) lists 1770 taxa, which, however, include many forms and varieties. For example, P. coccodes (Ach.) Nyl. has 17 forms and 10 varieties.

This Key includes 794 taxa, together with 316 synonyms or possible synonyms that are listed following the Key.

Please send any comments, corrections and additions to: [email protected] [email protected]

Chemistry

The compounds found in the genus Pertusaria fall into 8 categories:

Xanthones such as lichexanthone, 4,5-dichlorolichexanthone and thiophaninic acid.

Orcinol p-depsides such as lecanoric and gyrophoric acid.

Homologues of orcinol p-depsides such as perlatolic acid and its derivatives.

Orcinol depsones such as picrolichenic acid and its homologues.

-Orcinol m-depsides such as hypothamnolic and thamnolic acids.

-Orcinol p-depsides such as squamatic and barbatic acids.

-Orcinol depsidones such as stictic, norstictic, psoromic and protocetraric acids.

1 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

Aliphatic acids such as allopertusaric acid and lichesterinic acid.

In addition, some taxa lack lichen compounds.

Recently a number of Pertusaria species have been transferred to the genus Lepra Scop. (Scopoli 1777) by Hafellner & Türk (2016), Lendemer & Harris (2017), Buaruang et al. (2017) and Wei et al. (2017). This genus is characterised by disciform apothecia, asci with 1 (rarely 2) large or 8 small ascospores and the absence of chlorinated xanthones and orcinol p-depsides such as perlatolic acid and its derivatives. The published species are shown in the key with their authors; unpublished species are shown with Lepra only.

References:

Archer, A.W. (1997) A chemical and morphological arrangement of the lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 53:1-17.

Archer, A.W. (1995) A chemical and morphological arrangement of the lichen genus Pertusaria in Australia;: additional data and corrections. Mycotaxon 55:385-389.

Archer, A.W. (1997) The lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 69:5-249.

Buaruang, K; Boonpragob, K; Mongkolsuk, P; Sangvichien, E; Vongshewerat, K; Polyiam, W; Rangsiruji, A; Saipunkaew, W; Naksuwankul, K; Kalb, J; Parnmen, S; Kraichak, E; Phraphuchamnong, P; Meesim, S; Luangsuphabool, T; Nirongbut, P; Poengsungnoen, V; Duangphui, N; Sodamuk, M; Phokaeo, S; Molsil, M; Aptroot, A; Kalb, K; Lücking, R; Lumbsch, HT (2017): A new checklist of lichenised fungi occuring in Thailand. Mycokeys 23: 1-91.

Dibben, M.J. (1975) The Chemosystematics of the lichen genus Pertusaria in North America north of Mexico. Appendix II, Exotic species, pp. 492-519, Ph.D. thesis, Duke University.

Dibben, M.J. (1980) The Chemosystematics of the lichen genus Pertusaria in North America north of Mexico. Milwaukee Publications in Biology and Geology 5: 1-162.

Hafellner, J; Türk, R (2016): Die lichenisierten Pilze Österreich— eine neue Checkliste der Bisher nachgewiesen Taxa mit Angaben zu Verbreitung und Substratökologie. Stapfia 104(1):1-216.

Lendemer, JC; & Harris, JC (2017): Nomenclatural changes for North American members of the Variolaria- group necessitated by recognition of Lepra () Bryologist 120(2):183-190.

Hafellner, J; Türk, R (2016): Die lichenisierten Pilze Österreich— eine neue Checkliste der Bisher nachgewiesen Taxa mit Angaben zu Verbreitung und Substratökologie. Stapfia 104(1):1-216.

Hanko, B. (1983) Die Chemotypen der Flechtengattung in Europa. Bibliotheca Lichenologica 19:3-296.

Lendemer, JC; & Harris, JC (2017): Nomenclatural changes for North American members of the Variolaria- group necessitated by recognition of Lepra (Pertusariales) Bryologist 120(2):183-190.

Lumbsch, H.T. (1998) Taxonomic use of metabolic data in lichen-forming fungi. In: Frisvald, J.C., Bridge, P.D. & Arora, D.K (eds.) Chemical fungal : 345-387. Marcel Dekker, New York.

Lumbsch, H.T. & Nash, T.H (2001) Pertusaria in Nash, H.T., Ryan, B.D., Gries, C & Bungartz, F. Lichen Flora of the Greater Sonoran Desert Region 1: 341-357

Messuti, M.I. & Archer, A.W. (2009) ¿Cuántos taxones pueden incluirse teóricamente en el género Pertusaria? Glalia 2(1): 1-9.

Niebel-Lohmann, A. & Feuerer, T. (1992). Die Gattung Pertusaria DC (Lichenes) in Schleswig-Holstein: Anatomie, Morphologie Taxonomie und Verbreitung 1. Mitt. Inst. Allg. Bot. Hamburg. 24: 199-252.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

Oshio, M (1968) Taxonomical studies on the family Pertusariaceae of Japan. Journal of Science of the Hiroshima University, B(2), 12: 81-163.

Poelt, J. & Vezda, A. (1981). Pertusaria in Bestimmungsschlüssel europäischer Flechten. Ergänzungheft II. Bibliotheca Lichenologica 16: 1-390.

Scopoli, G.A. (1777) Introduct. ad Hist. Nat., W. Gerle, Prague

Wei, X; Schmitt, I; Hodkinson, B; Flakus, A; Kukwa, M; Divakar, K; Kirika, P; Otte, J; Meiser, A; Lumbsch, HT (2017) Circumscription of the genus Lepra, a recently resurrected genus to accommodate the "Variolaria"- Group of Pertusaria sensu lato (Pertusariales, ) PloS ONE 12(7): e0180284. https:/doi.org/10.137/journal.pone.0180284

Notes: The key uses apothecial structure and number and type of ascospores to place specimens in appropriate Groups (vide infra). Within the Groups chemical constituents, and size and arrangement of ascospores, are used to distinguish species. Synomyms and possible synonyms are indicated by the symbol  Only major chemical constituents are referred to and there remain species whose chemistry is unknown; these species are omitted until their chemistry is determined and published. Where the results of chemical tests (K, C, etc.) are reported and give a strong indication of the substance(s) present, these species are tentatively included.

Rock includes species growing on soil and plant débris, and mosses.

Disciform species include species where the apothecia is disciform and either sessile or raised in protuberances.

Sterile sorediate or isidiate species include species that may also occur as fertile specimens; sorediate or isidiate species which are always fertile are included in the appropriate fertile Group. Taxa which are reported to occur on both wood and rock are included twice.

Species with 2-3, or 2-4 ascospores per ascus are included in the 4-spored species; species with 6, 7 or 8 ascospores are inclcuded in 8-spored species.

Many images of tropical species of Pertusaria can be found on the web site: www.tropicallichens.net/ and bibliographic data on each species is available fom the Index Fungorum.

Group 1. Apothecia absent; isidia present; on rock Group 2. Apothecia absent; isidia present; on wood Group 3. Apothecia absent; soredia present; on rock Group 4. Apothecia absent; soredia present; on wood

Group 5. Apothecia present, verruciform; ascospores 1 per ascus; on rock Group 6. Apothecia present, verruciform; ascospores 1 per ascus; on wood Group 7. Apothecia present, disciform; ascospores 1 per ascus; on rock Group 8. Apothecia present, disciform; ascospores 1 per ascus; on wood

Group 9. Apothecia present, disciform; ascospores 2 per ascus; on wood Group 10 Apothecia, present disciform; ascospores 2 per ascus; on rock

Group 11. Apothecia present, verruciform; ascospores 2 per ascus, smooth; on rock Group 12. Apothecia present, verruciform; ascospores 2 per ascus, rough; on rock Group 13. Apothecia present, verruciform; ascospores 2 per ascus, smooth; on wood Group 14. Apothecia present, verruciform; ascospores 2 per ascus, rough; on wood

Group 15. Apothecia present, verruciform; ascospores 4 per ascus, smooth; on rock

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Group 16. Apothecia present, verruciform; ascospores 4 per ascus, rough; on rock Group 17. Apothecia present, verruciform; ascospores 4 per ascus, smooth; on wood Group 18. Apothecia present, verruciform; ascospores 4 per ascus, rough; on wood

Group 19. Apothecia present, disciform; ascospores 8 per ascus, on rock Group 20. Apothecia present, disciform; ascospores 8 per ascus, on wood Group 21. Apothecia present, verruciform; ascospores 8 per ascus, on rock Group 22. Apothecia present, verruciform; ascospores 8 per ascus, on wood

Sterile, lacking isidia, soredia and apothecia.

P. lichexanthonoverrucosa Aptoot & M. Cáceres; corticolous; lichexanthone & norstictic acid; SAm P. scepusiensis (Gyeln.) Erichs.; saxicolous; P-ve; Eur. P. shenandoensis Hale & Dibben; saxicolous; norstictic acid present; N.Am

Group 1. Apothecia absent; isidia present; on rock or plant débris

1. Xanthones present ...... 2 1a. Xanthones absent ...... 5

2. UV+ yellow; lichexanthone present; SAm...... P. subcorallina Nyl. 2a. UV+ orange or UV-ve; thiophaninic or thiophanic acid present ...... 3

3. UV-ve; thiophanic acid present; Qld ...... P. hiatensis A.W.Archer & Elix 3a. UV+ve; thiophaninic acid present ...... 4

4. Thiophaninic acid alone; UK, Sweden ...... P. flavocorallina Coppins & Muhr 4b. Thiophaninic acid and stictic acid present; Mediterranean region ...... P. rupicola (Fr.) Harm. var. coralloides (Anzi) Croz.

5. Thallus K+ red or yellow ...... 6 5a. Thallus K-ve ...... 12

6. K+ yellow; thamnolic or baeomycesic acid present ...... 7 6a. K+ red; norstictic or salazinic acid present ...... 8

7. Thamnolic acid present; Eurasia; also fertile with 2 spored asci ...... Lepra corallina (L.) Hafellner ...... [P. corallina (L.) Arn.] 7a. Baeomycesic and squamatic acids present; Aus...... P. (Lepra) nerrigensis A.W.Archer & Elix

8. Norstictic acid present ...... 9 8a. Salazinic acid present; Tasmania .. Lepra pseudodactylina (A.W.Archer) A.W.Archer & Elix [fertile in PNG;see Group 5] ...... [P. pseudodactylina A.W.Archer]

9. Lacking apothecia and spores ...... 10 9a. Sometimes fertile...... 11

10. East Asia ...... P. corallina var. minor (Yas. ex Räs.) Oshio

10a. Tasmania ...... Lepra dactylinella (Kantvilas & Elix) A.W.Archer & Elix ...... [P. dactylinella Kantvilas & Elix]

11b. N.Am, Eur; China; fertile in Eur...... P. pseudocorallina (Lilj.) Arnold 11c. Bipolar; also on wood; also 1/ascus 120-130 µm ...... P. coccodes (Ach.) Nyl.

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12(5a).Pd+ red; fumarprotocetraric or protocetraric acid present ...... 13 12a. Pd-ve ...... 17

13. Protocetraric acid present ...... 14 13a. Fumarprotocetraric acid present ...... 16

14. Protocetraric acid present ...... 15 14a. Protocetaric and hypothamnolic acids present; SAm...... P. pachythallina (Räs.) Messuti

15. Isidia to 1 mm tall; Antarctic; also fertile, 1 per ascus . Lepra corallophora (Vain.) Hafellner ...... [P. corallophora Vain.] 15a. Isidia coralloid, to 20 mm tall; SAm ...... Lepra acroscyphoides (Sipman) I. Schmitt, Hodkinson & Lumbsch ...... [P. acroscyphoides Sipman]

16. Isidia 1-3 mm tall; N. hemisphere; also fertile, 8 per ascus ...... P. oculata (Dickson) Th. Fr. 16a. Fumarprotocetraric and succinoprotocetraric acids present ...... P. dealbata (Ach.) Cromb.

17(12a). Confluentic or 2'-O-methylperlatolic acids present ...... 18 14a. Fatty acids present ...... 19

18. Confluentic acid present; Aus; Hafellner 15754a ...... Pertusaria sp. 18a. 2'-O-Methylperlatolic acid present; also fertile, 2 /ascus; Antarctic...... P. signyae Øvstedal

19. Isidia densely branched in upper parts; Antarctica ...... P. peudoculata Øvstedal 19a. Isidia to 1 mm tall; bipolar ...... P. isidiodes (Schaer.) Arnold

Group 2. Apothecia absent; isidia present; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 18

2. Lichexanthone or derivatives present ...... 3 2a. Norlichexanthone derivatives or thiophaninic acid present ...... 15

3. Lichexanthone present ...... 4 3a. Chlorolichexanthones present ...... 6

4. Lichexanthone present ...... 5 4a. Lichexanthone, 2-chlorolichexanthone and superlatolic acid present; Qld...... P. glebulosa A.W.Archer & Elix

5. Lichexanthone and salazinc acid present; PNG ...... P. parmatica A.W.Archer & Elix 5a. Lichexanthone and stictic acid; Thailand; ...... P. flavodigitata Jariang.

6. 2-Chlorolichexanthone, stictic and 2'-O-methyl- and 2-O-methylperlatolic acids present; ..... Australia ...... P. bagoensis Elix & A.W.Archer 6a. Polychlorolichexanthones present ...... 7

7. 4,5-Dichloro- and 2,4,5-trichlorolichexanthone present; Aus...... P. palumensis Elix & A.W.Archer 7a. 4,5-Dichlorolichexanthone present ...... 8

8. 4,5-Dichlorolichexanthone and depsides or depsidones present ...... 9 8a. 4,5-Dichlorolichexanthone and chiodectonic acid present; isidia red inside; Thailand ...... [Jariangprasert 4933.1] I am sure this is a Gassicurtia species – also in Australia“P. rufodigitata”

9. 4,5-Dichlorolichexanthone with various depsides; Aus...... P. georgeana and varieties 2-O-Methylperlatolicacid...... P. georgeana var. georgeana A.W.Archer & Elix

5 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

. 2-O-Methylperlatolic and 2-O-methylstenosporic acids ...... P. georgeana var. methylstenosporica A.W.Archer & Elix . 2,4-Di-O-methylolivetoric acid ...... P. georgeana var. goonooensis A.W.Archer & Elix . planaic acid ...... P. georgeana var. victoriana A.W.Archer & Elix . 2-O-Methylconfluentic acid- ...... P. georgeana var. occidentalis Elix & A.W.Archer 9a. 4,5-Dichlorolichexanthone with depsidones or other depsides ...... 10

10. 4,5-Dichlorolichexanthone and stictic acid present ...... 11 10a. 4,5-Dichlorolichexanthone and alectoronic acid present ...... 14

11. 4,5-Dichlorolichexanthone and stictic acid present ...... 12 11a. 4,5-Dichlorolichexanthone, stictic acid and depsides present ...... 13

12. Sterile; isidia 0.4-1 mm tall, 0.25-0.5 mm diam; Aus., PNG ...... P. montpittensis A.W.Archer 12a. Also fertile [Group 17]; isidia 0.1-0.5 mm tall; to 0.1 mm diam.; Eur ...... P. coronata (Ach.) Th.Fr.

13. 4,5-Dichlorolichexanthone, stictic and 2'-O-methylperlatolic acid acids present; Aus...... P. pilosula A.W.Archer & Elix 13. 4,5-Dichlorolichexanthone, stictic acid and methyl barbatate present; PNG ...... P. angabangensis A.W.Archer & Elix

14. Alectoronic acid present; Aus...... P. alectoronica Elix & A.W.Archer 14a. Alectoronic and thiophanic acids present; Aus ...... P. alectoronica var. thiophanica Kantvilas, Elix & A.W.Archer

15(2a).Thiophaninic and stictic acids present ...... 16 15a. Arthothelin or thiophanic acid present ...... 17

16 Aus & PNG ...... P. flavoisidiata A.W.Archer & Elix 16a. Eur..: rarely fertile, spores 8/ascus ...... P. flavida (DC) J.R. Laundon

17. Arthothelin and 6-O-methylarthothelin present; New Caledonia ...... P. simoneana A.W.Archer & Elix 17a. Thiophanic acid present; Qld...... P. lumbschii A.W.Archer & Elix

18 (1a).Lichesterinic acid present or lichen compounds absent ...... 19 18a. Depsidones, depsides or depside esters present ...... 20

19. Lichesterinic acid present; Aus...... P. (Lepra) ambigua A.W.Archer & Elix 19a. Lichen compounds absent; West Indies ...... P. griseola Vain.

20. Depsidones present [such as norstictic, protocetraric or psoromic acids] ...... 21 20a. Depsides or depside esters present [such as perlatolic acid derivatives] ...... 33

21. Stictic acid or derivatives present ...... 22 21a. Hypoprotocetraric, neotricone, norstictic, protocetraric, psoromic or salazinic acids present ...... 28

22. Stictic acid present ...... 23 22a. Hypostictic or norstictic acid present ...... 25

23. Stictic acid alone ...... 24 23a. Stictic and 2'-O-methylperlatolic acids present; Thailand, PNG ...... P. pilosula var. abditiva Jariang.

24. On wood; isidia 0.2-0.8 mm tall, 0.05-0.1 mm diam., Mauritius, Aus...... Lepra muricata( J.C. David) A.W.Archer & Elix ...... [P. muricata J.C. David]

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24a. On plant débris; isidia 0.5-1.5 mm tall, 0.15-0.25 mm diam.; NZ ...... P. (Lepra) dennistonensis Elix & A.W.Archer

25. Hypostictic and stictic acids present; Thailand ...... P. (Lepra) hypostictica Jariang. 25a. Norstictic acid present ...... 26

26. Isidia to 3 mm tall; Hawai; PNG ...... P. (Lepra) ramulifera H. Magn 26a. Isidia < 3mm tall...... 27

27. Isidia 0.1-0.25 mm tall; Aus., PNG ...... Lepra roseola (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. roseola A.W.Archer & Elix] 27a. Isidia 0.5-1.0 mm tall; bipolar; also on rock ...... P. (Lepra) coccodes (Ach.) Nyl.

28(21a). Protocetraric or hypoprotocetraric acids present ...... 29 28a. Neotricone, psoromic or salazinic and norstictic acids present ...... 31

29. Hypoprotocetraric acid present; PNG...... P. (Lepra) hypoprotocetrarica A.W.Archer & Elix 29a. Protocetraric acid present ...... 30

30. Protocetraric acid present; Aus. .. Lepra umbricola (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. umbricola A.W.Archer & Elix] 30a. Protocetraric and norstictic acids present; Aus...... Lepra wallamanensis (Elix & A.W.Archer) A.W.Archer & Elix ...... [P. wallamanensis Elix & A.W.Archer]

31(28a) Psoromic acid or neotricone present; ...... 32 31a. Norstictic and salazinic acids present; PNG; [HS 19405; CANB] Diorygma sp.? . Pertusaria sp.

32. Psoromic acid present; PNG ...... P. (Lepra) wauensis Elix & A.W.Archer 32a. Neotricone present; Aus...... Lepra) neotriconica (Elix & A.W.Archer) A.W.Archer & Elix ...... [P. neotriconica Elix & A.W.Archer]

33(20a).Methyl 3--hydroxy-4-O-demethylbarbatate present; PNG ...... P. mankiensis A.W.Archer & Elix 33a. Other depsides present ...... 34

34. Thamnolic acid present; Eur.; also fertile; 2 spores/ascus; isidia short and thick, ...... 0.1-0.3 mm diam ...... Lepra corallina (L.) Hafellner ...... [P. corallina (L.) Arn.] Isidia very thin, 0.03-0.05 mm diam.; Aus...... Lepra tricosa (Elix & A.W.Archer) A.W.Archer & Elix ...... [P. tricosa Elix & A.W.Archer] 34a. 2'-O-Methylperlatolic, barbatic or divaricatic acids or derivatives or atranorin present ...... 35

35. 2'-O-Methylperlatolic acid present; PNG, Aus...... P. burburana Elix & A.W.Archer 35a. Barbatic or divaricatic acid or atranorin and bourgeanic acid present ...... 36

36. Atranorin and bourgeanic acid present ...... P. ochrodigitula A.W.Archer & Elix 36a. Barbatic or divaricatic acid present ...... 37

37. Barbatic acid present; Aus...... Lepra barbatica (A.W.Archer & Elix) I. Schmitt, Hodkinson & Lumbsch ...... [P. barbatica A.W.Archer & Elix] 37a. Divaricatic acid ± stictic acid present; PNG, Aus ...... P. labuensis A.W.Archer & Elix

Group 3. Apothecia absent; soredia present; on rock or plant débris.

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1. Xanthones present ...... 2 1a. Xanthones absent ...... 17

2. Lichexanthone or chlorolichexanthones present ...... 3 2a. Thiophaninic acid or thiophanic acids present ...... 11

3. Chlorinated lichexanthones present ...... 4 3a. Lichexanthone present ...... 5

4. 2,4- and 2,5-Dichlorolichexanthone, 2,4,5-trichlorolichexanthone and stictic acid present ..... Australia; PNG ...... P. puffina A.W.Archer & Elix 4a. 4,5-Dichlorolichexanthone and stictic acid present; PNG; also corticolous ...... P. balekensis A.W.Archer & Elix

5. Lichexanthone and depsides present ...... 6 5a. Lichexanthone and picrolichenic acid present ...... 8

6. Lichexanthone and methyl 2'-O-methylmicrophyllinate present; Aus...... P. variabilis Elix & A.W.Archer ...... [Transfer to Lepra not accepted; depside present.] 6a. Lichexanthone and hypothamnolic or haemathamnolic acid present ...... 7

7. Hypothamnolic acid present ...... Lepra amaroides (H. Magn.) I. Schmitt, Hodkinson & Lumbsch ...... [P. amaroides H. Magn.] Hypothamnolic and barbatic acids present; Uruguay; 7031 ...... Pertusaria sp. 7a. Haemathamnolic acid present; Brazil ...... P. xantholeucoides Müll. Arg.

8. Lichexanthone and picrolichenic acid alone or with superpicrolichenic acid ...... 9 8a. Lichexanthone, picrolichenic acid and -orcinol m-depsides...... 10

9. Lichexanthone and picrolichenic acid present; Aus. and S.Am...... [rarely with additional norstictic acid] ...... Lepra subventosa var. deficiens (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. subventosa var. deficiens A.W.Archer & Elix] 9a. Lichexanthone, picrolichenic and superpicrolichenic acids present; Norfolk Island, Aus...... Lepra verdonii (A.W.Archer) I. Schmitt, Hodkinson & Lumbsch ...... [P. verdonii A.W.Archer]

10. Lichexanthone, picrolichenic and thamnolic acid present; S.Am, Aus. and NZ; ...... very rarely fertile ...... P. subventosa var. subventosa Malme 10a. Lichexanthone, picrolichenic and hypothamnolic acid present; Aus., PNG ...... Lepra subventosa var. hypothamnolica (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. subventosa var. hypothamnolica A.W.Archer & Elix] [Lichexanthone, picrolichenic and barbatic acid present; S.Am; Ahti 31.i.1989 (H) ...... Pertusaria sp. ]

11(2a). Thiophanic acid and arthothelin present; on plant débris; Qld...... P. "finniganensis" 11a. Thiophaninic acid present ...... 12

12. Thiophaninic acid, confluentic and planaic acids present; N.Am.; ± fertile ...... P. tejocotensis de Lesd. 12a. Thiophaninic acid and depsidones present ...... 13

13. Thiophaninic acid and norstictic acid (major) and stictic acid (minor) present; Aus...... P. rogersii A.W.Archer & Elix 13a. Thiophaninic acid and stictic acid present ...... 12

14. In Australia ...... 15 14a. Elsewhere ...... 16

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15. Thallus bright yellow; hypostictic acid absent; E Australia, NZ & PNG; ± fertile ...... P. xanthoplaca Müll. Arg. 15a. Thallus dull yellow; hypostictic acid present; NW Aus...... P. remota A.W.Archer

16. Eur., Asia, Africa ...... P. amarescens Nyl. Japan ...... P. leucosoroides Nyl. 16a. N.Am.; ± fertile ...... P. flavicunda Nyl.

17. Lichen compounds absent ...... 18 17a. Lichen compounds present ...... 19

18. Canada ...... P. pruinifera Erichs. 18a. Philippines ...... P. subvaginata var. orientalis Räs.

19 Picrolichenic acid or depsides present [such as lecanoric or squamatic acids]...... 20 19a. Depsidones present [such as norstictic or protocetraric acids and derivatives] ...... 24

20. Picrolichenic acid present ...... P. amara (Ach.) Nyl. Additional protocetraric acid present; Eur...... P. amara var. flotowiana (Flörke) Erichs. 20a. Depsides present ...... 21

21. Lecanoric acid present ...... 22 21a. Other depsides present ...... 23

22. Lecanoric acid present, variolaric acid absent; Philippines ...... P. subvaginata var. orientalis Räs. 22a. Lecanoric and variolaric acids present; Europe; ± fertile ...... P. lactea (L.) Arnold

23. Squamatic and baeomycesic acids present; Aus...... Lepra nerrigensis (A.W.Archer & Elix) I. Schmitt, Hodkinson & Lumbsch ...... [P. nerrigensis(A.W.Archer & Elix] 23a. 2'-O-Methylisohyperlatolic and 2'-O-methylsuperlatolic acids present; Aus...... P. salebrosa A.W.Archer & Elix

24(19a).Norstictic or stictic acid present ...... 25 24a. Protocetraric or fumarprotocetraric acid present ...... 28

25. Stictic acid present; soralia ochre coloured; S. Georgia ...... Pertusaria sp. A Øvstedal 25a. Norstictic acid present ...... 26

26. Norstictic acid alone ...... 27 26a. Norstictic and picrolichenic acids present; Teneriffe .... Lepra teneriffensis (Vain.) Hafellner ...... [P. teneriffensis Vain.

27. Thallus off-white; Europe, China, N.Am, ...... Lepra excludens (Nyl.) Hafellner ...... [P. excludens Nyl.] 27a. Thallus olive green; Aus., Lord Howe Is ...... Lepra miniatescens (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. miniatescens A.W.Archer & Elix]

28. Protocetraric acid present; Europe, N.Afr., Japan, Korea .... Lepra leucosora (Nyl.) Hafellner ...... [P. leucosora Nyl.] Canary Islands ...... P. inconveniens Vain. 28a. Fumarprotocetraric acid present ...... 29

29a. Atranorin present; soralia off-white; subtropical; Aus...... Lepra sordida( A.W.Archer) A.W.Archer & Elix ...... [P. sordida A.W.Archer]

9 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

29a. Atranorin absent; ± succinprotocetraric acid present; Eur ...... 30

30. Thallus on limestone ...... P. gypsicola Erichs. 30b. Thallus on siliceous rocks ...... 31

31. Thallus with flattened, white soralia...... Lepra aspergilla (Ach.) Hafellner ...... [P. aspergilla (Ach.) J.R. Laundon] 31a. Thallus with sorediate papillae ...... Lepra mammosa (Harm.) Hafellner ...... [P. mammosa Harm.]

Group 4. Apothecia absent; soredia present; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 15

2. Arthothelin, thiophaninic acid or thuringione present ...... 3 2a. Lichexanthone or 4,5-dichlorolichexanthone present ...... 5

3. Arthothelin present ...... 4 3a. Thiophaninic acid and depsidones present ...... 8

4. Arthothelin, thiophaninic acid and barbatic acid present;Thailand . P. flavosorediata Jariang. 4a. Arthothelin and thuringone present; Australia ...... P. flavopunctata A.W.Archer & Elix

5(2a). 4,5-Dichlorolichexanthone present ...... 6 5a Lichexanthone present alone orwith depsone, depsidone, depside or fatty acid ...... 9

6. 4,5-Dichlorolichexanthone present alone; PNG; also saxicolous ...... P. balekensis A.W.Archer & Elix 6a. 4,5-Dichlorolichexanthone with depside or depsidone present ...... 7

7. 4,5-Dichlorolichexanthone and stictic acid present; N.Am ...... P. expolita Harris 7a. 4,5-Dichlorolichexanthone and planaic acid present; Aus ...... P. puttyensis A.W.Archer & Elix

8. .Thiophaninic acid and stictic acid present; Aus...... P. maritima A.W.Archer & Elix 8a. Thiophaninic acid and norstictic acid present; N.Am...... P. azulensis de Lesd.

9. Lichexanthone only present; SAm ...... P. lichexanthofarinosa Aptroot & M. Cáceres 9a. Lichexanthone with other compounds ...... 10

10. Lichexanthone, picrolichenic and superpicrolichenic acids present; Norfolk Island, Aus...... P. (Lepra) verdonii (A.W.Archer) I. Schmitt, Hodkinson & Lumbsch 10a. Lichexanthone with other compounds present ...... 11

11. Lichexanthone and confluentic or thamnolic acid present ...... 12 11a. Lichexanthone and stictic or dihydropertusaric acid present ...... 14

12. Lichexanthone and confluentic acid present; Aus., Thailand ...... P..confluentica Jariang. 12a. Lichexanthone and thamnolic acid present...... 13

13. Soralia flattened, inconspicuous; Sri Lanka ...... Lepra leucosorodes (Nyl.) I.Schmitt, Hodkinson & Lumbsch ...... [P. leucosorodes Nyl.] 13a. Soralia globular, conspicuous; Aus., PNG ...... Lepra scaberula (A.W.Archer) I. Schmitt, Hodkinson & Lumbsch ...... [P. scaberula A.W.Archer]

10

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

14. Lichexanthone and stictic acid present; Hawaii ...... Lepra oahuensis (H. Magn.) A.W.Archer & Elix ...... [P. oahuensis H. Magn.] 14a. Lichexanthone and dihydropertusaric acid; Fiji ...... P. cisalbescens Elix & A.W.Archer

15(1a). Fatty acids present ...... 16 15a. Depsides or depsidones present or lichen compounds absent ...... 19

16. Lichen compounds absent; Norfolk Island ...... P. heinarii A.W.Archer & Elix 16a. Fatty acids present ...... 17

17. Allopertusaric acid present; Northern Hemisphere ...... Lepra albescens (Huds.) Hafellner ...... [P. albescens (Huds.) Choisy & Werner] 17a. Lichesterinic acid or isomyelochroic acid present ...... 18

18. Lichesterinic acid present; Central Africa ...... P. krogiae A.W.Archer et al. 18a.. Isomyelochroic acid present ...... Lepra ophthalmiza (Nyl) Hafellner ...... [P. ophthalmiza (Nyl.) Nyl.]

19(15a).Depsidones present ...... 20 19a. Depsides or picrolichenic acid present or lichen compounds absent ...... 27

20. Norstictic, stictic or fumarprotocetraric acid present ...... 21 20a. Psoromic acid present; Aus., NZ, PNG ...... Lepra psoromica (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. psoromica A.W.Archer & Elix]

21. Stictic acid present; Aus...... Lepra albopunctata (A.W.Archer & Elix) A.W.Archer & Elix ...... [P. albopunctata A.W.Archer & Elix] 21a. Norstictic or fumarprotocetraric acid present ...... 22

22. Fumarprotocetraric acid present; Eur, N.Am; also fertile ...... Lepra borealis (Erichs.) I.Schmitt, Hodkinson & Lumbsch ...... [P. borealis Erichs.] 22a. Norstictic acid present ...... 23

23. Australia; soralia white or pale olive-green ...... 24 23a. Elswhere ...... 26

24. Soralia white; ...... 25 24a. Soralia pale olive-green; soralia covering thallus surface, ± sekikaic acid ...... P. scabrida A.W.Archer & Elix

25. Soralia 0.5-1 mm diam., well-defined; stictic acid absent...... Lepra erythrella (Müll. Arg.) I.SChmitt, Hodkinson & Lumbsch ...... [P. erythrella Müll. Arg.] 25a. Soralia to 2 mm diam, ill-defined; with additional stictic acid; ...... P." alloerythrella"

26. India; soralia ochraceous to ochraceous red, 0.5-2 mm diam...... P. colorata Awasthi & Srivastava 26a.. West Indies; K+ red; soralia 0.4-2 mm diam...... P. torulosa Vain.

27(19a). Lichen compounds absent; Philippines ...... P. amara var. philippinensis Räs.

27a. Depsides or depsone present ...... 28

28. Thamnolic or 2'-O-methylperlatolic or hypothamnolic acid or atranorin present ...... 29 28a Lecanoric or picrolichenic acids present ...... 33

29. 2'-O-Methylperlatolic acid or thamnolic acid present: ...... 30

11 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

29a. Hypothamnolic acid or atranorin present ...... 32

30. 2'-O-Methylperlatolic acid present; Thailand ...... P. uttaraditensis Jariang. 30a. Thamnolic acid present ...... 31

31. Thallus surface sorediate; Japan, Aus...... Lepra violacea (Oshio) I. Schmitt, Hodkinson & Lumbsch ...... [P. violacea Oshio] 31a. Thallus surface granular; N.Am ...... Lepra pustulata (Brodo & W.L. Culb.) Lendemer & R.C. Harris ...... [Haematomma pustulata Brodo & W.L. Culb.] 32. Hypothamnolic and cryptothamnolic acids present...... Lepra subviolacea (Q. Ren) I. Schmitt, Hodkinson & Lumbsch ...... [P. subviolacea Q. Ren] 32a.. Atranorin present; soralia flat; Hawaii and Aus...... P. sellingii H.Magn.

33.(28a) Lecanoric and variolaric acids present ...... P. hemisphaerica (Flörke) Erichs. 33a. Picrolichenic acid present ...... 34

34. Picrolichenic acid only present; K-, C-, KC+ reddish; Hawaii ...... P. molokaiensis H. Magn. 34a. Picrolichenic and other acids present ...... 35

35. Picrolichenic ± protocetraric acids present; ± fertile ...... Lepra amara (Ach.) Hafellner ...... [P. amara (Ach.) Nyl.] 35a. Picrolichenic and subpicrolichenic acids present; PNG ...... Lepra buloloensis (A.W.Archer, Elix & Streimann) Schmitt & Lumbsch ...... [P. buloloensis A.W.Archer, Elix & Streimann]

Group 5. Apothecia present, verruciform; ascospores 1 per ascus; on rock

1. 5-O-Methylhiasic acid present; spores 130-20 µm; Aus .... P. flavoexpansa Kantvilas & Elix 1a. Norstictic or protocetraric or salazinic acid present ...... 2

2. Norstictic or protocetraric acid present ...... 3 2a. Salazinic acid present; spores 180-200 µm long ...... Lepra pseudodactylina (A.W.Archer) A.W.Archer & Elix [sterile in Tas; see Group 1] ...... [P. pseudodactylina A.W.Archer]

3. Norstictic acid present; spores 170-395 µm long; NZ ...... P. otagoana Galloway 3a. Protocetraric acid present; spores ca. 140 µ long; Antarctic; also sterile ...... P. corallophora Vain.

Group 6. Apothecia present, verruciform; ascospores 1 per ascus; on wood

1. Xanthones absent; protocetraric or fumarprotocetraric acid present ...... 2 1a. Xanthones present; protocetraric or fumarprotocetraric acid absent ...... 3

2. Protocetraric acid present; thallus not isidiate; spores 220-225 µm long; Indonesia ...... P. leeuwenii Zahlbr. 2a. Fumarprotocetraric acid present; thallus isidiate; spores 160-280 µm long; Greece ...... Lepra graeca (Erichs.) Hafellner ...... [P. graeca Erichs.]

3. Thiophaninic acid present ...... 4 3a. Lichexanthone or 4,5-dichlorolichexanthone present ...... 7

4. Thiophaninic acid only present; spores 142-187 µm long; ostioles black, disciform; Canary Is, ...... P. calderae Hern.-Padr et al.,

12

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

4a. Thiophaninic acid and 2'-O-methylperlatolic or confluentic acid present ...... 5

5. Confluentic acid present; spores 150-250 µm long; Greece ...... P. corinthiaca Erichs. 5a. 2'-O-Methylperlatolic acid present; spores 120-220 µm long ...... 6

6. Spores 180-220 µm long; Eur ...... P. caesioalba (Flot.) Nyl. 6a. Spores 120-150µm long; ± planaic acid; Spain, Turkey .... P. paramerae A.Crespo & Vĕzda

7. Lichexanthone present; spores 170-225 µm long; S.Am. [also 2/ascus] ...... P. tetrathalamia var major Müll.Arg. 7a. 4,5-Dichlorolichexanthone and stictic acid present; spores rarely 1 per ascus, 262 x 45 µm, . usually 4 per ascus (q.v.); West Indies ...... P. ascidioides Vain.

Group 7. Apothecia present, disciform; ascospores 1 per ascus; on rock or moss or plant debris

1. Thallus isidiate ...... 2 1a Thallus not isidiate ...... 10

2. Fatty acids or 5-O-methylhiascic acid present; fumarprotocetraric acid absent ...... 3 2a. Fumarprotocetraric acid present ...... 7

3. Fatty acids present ...... 4 3a. 5-O-Methylhiascic acid present; spores 150-200 µm long; on soil or rock; Tas ...... [pseudodisciform; see Group 5] ...... P. flavoexpansa Kantvilas & Elix

4. On rock ...... 5 4a. On moss or plant débris ...... 6

5. Spores 200 µm long; Ireland ...... P. polythecia (Taylor) Erichs. 5a. Spores 122-183 µm long; Eurasia ...... Lepra ocellata (Körb.) Hafellner ...... [P. ocellata Wallr. ex Körb.]

6. On moss; spores 80-165 µm long; Ireland ...... P. hutchinsiae (Borrer) Leight. 6a. On plant débris; spores 80-165 µm long; N. hemisphere ...... Lepra panyrga (Ach.) Hafellner ...... [P. panyrga (Ach.) Massal.]

7 Isidia becoming sorediate; spores 180-200 µm long; France ...... Lepra mammosa (Harm.) Hafellner ...... [P. mammosa Harm.] 7a. Isidia well-defined ...... 8

8. Apothecia terminal in isidia; spores 100-270 µm long; circumpolar ...... Lepra dactylina (Ach.) Hafellner ...... [P. dactylina (Ach.) Nyl.] 8a. Apothecia not terminal in isidia ...... 9

9. Isidia short how many mm, blunt, crowded; ?spores; Eur ... Lepra stalactiza (Nyl.) Hafellner ...... [P. stalactiza Nyl.] 9a. Isidia tall how tall?, scattered; spores 115-140 µm long; N.Am ...... Lepra andersoniae (Lendemer) Lendemer & R.C. Harris ...... [P. andersoniae Lendemer]

10(1a). Lichexanthone present ...... 11 10a. Lichexanthone absent ...... 16

11. Picrolichenic acid present ...... 12 11a Picrolichenic acid absent ...... 13

13 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

12. Lichexanthone and picrolichenic acid present; spores 125-180 µm long; N.Am, S.Am ...... Lepra ventosa (Malme) Lendemer & R.C. Harris ...... [P. ventosa Malme] 12a. Lichexanthone, picrolichenic and thamnolic acids present; spores 120-160 µm long, rare; .... Aus., S.Am ...... Lepra subventosa (Malme) Schmitt & Lumbsch ...... [P. subventosa Malme]

13. Hypothamnolic acid present; spores 110-160 µm long; S.Am ...... P. hossei (Räs.) A.W.Archer & Osorio 13a. Hypothamnolic acid absent ...... 14

14. Lecanoric acid present; spores 170-220 µm long; Juan Fernandez ... P. skottsbergii Zahlbr...... [also on wood] 14a. Lecanoric acid absent ...... 15

15. Haemathamnolic acid present; spores not reported; Brazil ...... Lepra xantholeucoides (Müll.Arg.) I. Schmitt, A.W.Archer & Lumbsch ...... [P. xantholeucoides Müll.Arg.] 15a. Squamatic acid present; spores 100-130 µm long; Socotra ...... Lepra xantholeuca (Müll. Arg.) I. Schmitt, A.W.Archer & Lumbsch ...... [P. xantholeuca Müll. Arg.]

16(10a). Lichen compounds absent; spores 90-140 µm long; Lapland ...... P. tornensis H.Magn. 16a. Lichen compounds present ...... 17

17. Spores 150-180 µm; K-ve, C-ve [2 unidentified compounds, hplc] Mexico ...... P. velata f. liebmannii Vain. 17a. Other compounds present ...... 18

18. Norstictic acid present ...... 19 18a. Norstictic acid absent ...... 21

19. Norstictic acid only present; spores 100-300 µm long; Eur ...... Lepra monogona (Nyl.) Hafellner ...... [P. monogona Nyl.] 19a. Norstictic and picrolichenic acids present ...... 20

20. Spores 150-160 µm long [usually 2 per ascus]; disc dull fawn to brown; S.Am...... Lepra alterimosa (Darb.) I. Schmitt, Hodkinson & Lumbsch ...... [P. alterimosa Darb.] 20a. Spores 100-150 µm long; disc black, white pruinose; Eur...... P. ludovicae Werner

21(18a). Picrolichenic and protocetraric acids present; spores 180-250 µm long; [rarely on wood] ...... Lepra melanochlora (DC) Hafellner ...... [P. melanochlora (DC) Nyl.] 21a. Picrolichenic acid absent ...... 22

22. Thamnolic, cryptothamnolic or hypothamnolic acids present ...... 23 22a. Lecanoric, protocetraric or gyrophoric acids present ...... 25

23. Thamnolic acid present ...... 24 23a. Cryptothamnolic and hypothamnolic acids present; spores 153-204 µm long; China; ...... [also on wood] ...... P. leptospora Nitschke ex Lahm

24. Spores 95-175 µm long; S.Afr...... Lepra wawreana (A.Massal.) I. Schmitt, Hodkinson & Lumbsch ...... [P. wawreana A.Massal.] 24a. Spores "ca.200 µm long" [fide Müll. Arg.]; Eur...... P. clementiana Müll. Arg.

14

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

25. Protocetraric acid present; ...... 26 25a. Lecanoric or gyrophoric acid present ...... 29

26. Southern Hemisphere ...... 27 26a. Northern Hemisphere ...... 28

27. Spores 145-185 µm long; S.Am,, NZ ...... Lepra macloviana (Müll. Arg.) I.Schmitt, Hodkinson & Lumbsch ...... [P. macloviana Müll. Arg.] 27a.. Spores 195-280 µm long; Tasmania ...... Lepra parathalassica (Kantvilas & Elix) A.W.Archer& Elix ...... [P. parathalassica Kantvilas & Elix]

28. Spores 135-240 µm long; Arctic Russia ...... P. cribellata Branth 28a. Spores 160-180 µm; China ...... P. setschwanica Zahlbr.

29. Lecanoric acid present ...... 30 29a. Gyrophoric acid present ...... 32 . 30. Lecanoric and variolaric acids present; spores 180-240 µm long; Eur ...... P. lactea (L.) Arn. 30a. Lecanoric acid only present ...... 31

31. Spores 100-200(-275) µm long; NZ ...... P. obvelata Nyl. 31a. Spores 200-270 µm long; France ...... P. conglobata (Ach.) Fr

32. Spores 135-150 µm long; S.Am ...... P. microcarpa Nyl. 32a. Spores 150-230 µm long; circumarctic ...... P. bryontha (Ach.) Nyl.

Group 8. Apothecia present, disciform; ascospores 1 per ascus; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 14

2. Chloroxanthones present ...... 3 2a. Lichexanthone present ...... 4

3. 5,7-Dichloro-3-O-methylnorlichexanthone present; spores 125-165 µm long; NZ ...... P. glaucomopsis Nyl. 3a. 2-Chlorolichexanthone and 2'-O-methylperlatolic acid present; spores 100-130 µm long; NZ ...... P. circumcincta Stirt. 

4. Lichexanthone only present; spores 100-150 µm long; PNG ...... P. asterella Aptroot 4a. Lichexanthone and picrolichenic acid or depsides present ...... 5

5. Lichexanthone and picrolichenic acid present ...... 6 5a. Lichexanthone and depsides present ...... 7

6. Spores 160-195 µm long; picrolichenic acid present; PNG ...... P. myola A.W.Archer & Elix 6a. Spores 135-175 µm long; picrolichenic acid and homologues present; Aus...... Lepra clarkeana (A.W.Archer) I.Schmitt, Hodkinson & Lumbsch ...... [P. clarkeana A.W.Archer]

7. Lecanoric acid present; spores 110-175 µm long [120-315 µm (Dibben)]; cosmopolitan ...... Varicellaria velata (Turner) I. Schmit & Lumbsch ...... [P. velata (Turner) Nyl.] 7a. Baeomycesic, thamnolic, hypothamnolic or haemathamnolic acid present ...... 8

8. Baeomycesic or thamnolic acid present ...... 9

15 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

8a. Hypothamnolic or haemathamnolic acid present ...... 10

9. Baeomycesic acid present; spores 68-198 µm long; N.Am ...... Lepra floridana (Dibben) Lendemer & R.C. Harris ...... [P. floridana Dibben] Baeomycesic and hypothamnolic acids present; spores 120-150 µm long; Brazil ...... P. (Lepra ) elizabethae A.W.Archer & Elix 9a. Thamnolic acid present; spores 100-130 µm long; Aus ...... Lepra miscella (A.W.Archer) I. Schmitt, Hodkinson & Lumbsch ...... [P. miscella A.W.Archer]

10. Hypothamnolic acid present ...... 11 10a. Haemathamnolic acid present ...... 13

11. Spores 115-150 µm long; Africa ...... Lepra tropica (Vain.) Lendemer & R.C. Harris ...... [P. tropica Vain.] 11a. Spores >150 µm long ...... 12

12 Spores 153-204 µm; China; [Pd+ yellow!]; also on rock ..... P. leptospora Nitschke ex Lahm 12a. Spores 174-210 µm long; with additional salazinic acid; China ...... Lepra paratropica (Ren) I. Schmitt, Hodkinson & Lumbsch ...... [P. paratropica Ren]

13. Spores 100-150 µm long; circumtropical ...... June 12. 2017 ...... Lepra commutata (Müll.Arg.) Lendermer & R.C. Harris July 11. 2017 ...... [Lepra commutata (Müll.Arg.) I. Schmitt, Hodkinson & Lumbsch] ...... [P. commutata Müll.Arg.] 13a. Spores 150-170 µm long; S.Am...... Lepra superans (Müll.Arg.) I. Schmitt, Hodkinson & Lumbsch ...... [P. superans Müll.Arg.]

14(1a). Fatty acids present; spores 100-200 µm long; N Eur & N.Am ...... P. panyrga (Ach.) Massal. 14a. Depsides, depsidones or picrolichenic acid present ...... 15

15. Depsidones [fumarprotocetraric, physodalic, protocetraric or norstictic acids] present ...... 16 15a. Depsides [lecanoric, squamatic, barbatic, thamnolic, haemathamolic or hypothamnolic ...... acids present] or picrolichenic acid present ...... 27

16. Fumarprotocetraric or protocetraric accid present ...... 17 16a. Physodalic or norstictic acid acid present ...... 21

17. Fumarprotocetraric acid present ...... 18 17a. Protocetraric acid present ...... 19

18. Spores 110-160 µm long; N.Am, Eur ...... Lepra multipunctoides (Dibben) Lendemer & R.C. Harris ...... [P. multipunctoides Dibben] 18a. Spores145-198 µm long; China ...... P. multipuncta Nyl. var. colorata Zahlbr.

19. Protocetraric acid and thamnolic acids present; spores 125-180 µm long; China ...... P. huangshanensis S.Yu & J.Wu ex Q. Ren 19a. Protocetraric acid present ...... 20

20. Spores 210-250 µm long; China ...... Lepra sejilaensis (Q. Ren) I. Schmitt, Hodkinson & Lumbsch ...... [P. sejilaensis Q. Ren] 20a. Spores 170-180 µm long; Aus...... Lepra lacericans (A.W.Archer) A.W.Archer & Elix ...... [P. lacericans A.W.Archer]

16

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

21. Physodalic acid present; spores 90-170 µm long [-242 µm, China]; Korea, Eur...... Lepra multipuncta (Turner) Hafellner ...... [P. multipuncta (Turner) Nyl.] 21a. Norstictic acid present ...... 22

22. Norstictic and salazinic acids present; spores 120-150 µm long; S.Afr...... P. casta Zahlbr. 22a. Norstictic acid present; salazinic acid absent ...... 23

23. Spores 200-300 µm long; thallus sorediate; Eur...... P. monogoniza Nyl. 23a. Spores <200 µm long ...... 24

24. Spores 150-175 µm long; disc white pruinose; apothecia pustulate; Aus, PNG ...... Lepra sublacerans (A.W.Archer) A.W.Archer & Elix ...... [P. sublacerans A.W.Archer] 24a. Spores < 160 µm long ...... 25

25. Discs black, epruinose; spores 100-160 µm long; Japan; China, Korea ...... P. submultipuncta Nyl. 25a. Spores 30-150 µm long ...... 26

26. Discs black or pink, pruinose; spores 70-150 µm long; Eur and N.Am...... Lepra waghornei (Hult.) Lendemer & R.C. Harris ...... [P. waghornei Hult.] 26a. Discs black; spores 30-80 µm long; Finland ...... P. atropallida Vain.

27(15a). Lecanoric or picrolichenic acid present ...... 28 27a. Squamatic, barbatic, thamnolic, haemathamolic or hypothamnolic acids present ...... 35

28. Lecanoric acid present ...... 29 28a. Picrolichenic ± protocetraric acid present ...... 31

29. South Africa; spores 125-160 µm long; ...... P. subvelatula Vain. 29a. Hawaii ...... 30

30. Spores 300-325 µm; ...... P. contraria H.Magn. 30a. Spores 134-236 µm; ...... P. hawaiiensis Erichs.

31. Spores 130-150µm long; ± protocetraric acid; also saxicolous and/or sterile ...... Lepra amara (Ach.) Hafellner ...... [P. amara (Ach.) Nyl.] 31a. Spores ca. >150 µm long ...... 32

32. Atranorin present ...... 33 32a. Atranorin absent ...... 34

33. Spores 150-170 µm long; Aus; apothecia dish-shaped...... Lepra patellifera (A.W.Archer) Schmitt & Lumbsch ...... [P. patellifera A.W.Archer] 33a. Spores 137-180 µm long; S.Am; apothecia hemispherical.P. grassiae Messuti & A.W.Archer

34. Epithecia yellow; spores 150-200 µm; NZ ...... Lepra flavovelata (Elix & Malcolm) I.Schmitt, Hodkinson & Lumbsch ...... [P. flavovelata Elix & Malcolm] 34a. Epithecia not yellow; spores 170-225 µm; Aus, PNG, Brazil, Fiji, Sri Lanka ...... Lepra lacerans (Müll.Arg.) I. Schmitt, Hodkinson & Lumbsch ...... [P. lacerans Müll.Arg.]

35(27a). Squamatic or barbatic acids present ...... 36 35a. Thamnolic, haemathamolic or hypothamnolic acids present ...... 37

17 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

36. Squamatic acid present; spores 112-140 µm long; Indonesia...... P. gedehana Zahlbr. 36a. Barbatic acid present; spores 200-24 µm long; NZ ...... Lepra wirthii (Elix & A.W.Archer) I. Schmitt, Hodkinson & Lumbsch ...... [Pertusaria wirthii Elix & A.W.Archer]

37. Thallus Pd-ve, K+ violet; hypothamnolic acid present; spores 140-170 µm long ...... Aus., NZ ...... Lepra novaezelandiae (Szatala) I. Schmitt, A.W.Archer & Lumbsch ...... [P. novaezelandiae Szatala] 37a. Thallus Pd+ yellow, K+ yellow ...... 38

38. Haemathamnolic acid present ...... 39 38a. Thamnolic acid present ...... 41

39. Spores 105-150 µm long; S.Am; Japan ...... P. aggregata Müll.Arg. 39a. Spores140-200 µm long ...... 40

40. Disc pale; spores 140-200 µm long; S.Am., Korea ...... Lepra variolosa (Kremp.) I. Schmitt, A.W.Archer & Lumbsch ...... [P. variolosa (Kremp.) Vain.] 40a. Disc black; spores 162-190 µm long; S,Am...... Lepra ornatula (Müll.Arg.) I. SChmitt, Hodkinson & Lumbsch ...... [P. ornatula Müll.Arg.]

41. Spores usually < 150 µm long ...... 42 41a. Spores usually ca. >150 µm long ...... 44

42. Thallus esorediate; disc black, pruinose; spores 112-132 µm long PNG ...... P. yupna A.W.Archer & Elix 42a. Thallus sorediate; soredia yellow or white ...... 43

43. Spores 100-150 µm long, soredia yellow; Lappland ...... P. efflorescens Vain. 43a. Spores 100-140 µm long; soredia white; Japan ...... P. variolina Nyl.

44. Disc orange, pruinose; spores 150-185 µm long; NZ ...... Lepra scutellifera (A.W.Archer & Elix) I. Schmitt, Hodkinson & Lumbsch ...... [P. scutellifera A.W.Archer & Elix] 44a. Disc black, epruinose; spores 140-230 µm long; Japan ...... P. nigrodisca Oshio

Group 9. Apothecia present, disciform; ascospores 2 per ascus; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 4

2. 4,5-Dichlorolichexanthone and perlatolic acid derivs. present; N.Am...... P. sulcata Dibben 2a. Thiophaninic acid present ...... 3

3. Thiophaninic acid only present; spores 125-182 µm long; Canary Islands ...... P. aceroae C.Hdez.-Padr. et al. 3a. Thiophaninic and norstictic acids present; spores 90-155 µm long; N.Am. P. lecanina Tuck.

4. Lichen compounds absent ...... 5 4a. Lichen compounds present ...... 7

5. N. hemisphere, spores 70-115 µm long; ?boreal ...... P. stenhamerii Helb. 5a. North and South America ...... 6

6. S. Am.; spores 75-90 µm long; epithecia K-ve ...... P. kalbii A.W.Archer & Messuti 6a. N.Am.; spores 80-110 µm long; epithecia K+ violet; ...... P. rhexostoma Nyl. ex Hue

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

7. Depsidones [protocetraric, norstictic or salazinic acids] present ...... 8 7a. Depsides [2-O-methylconfluentic, gyrophoric, lecanoric, planaic, thamnolic acids] present ...... 14

8. Protocetraric acid present; spores 70-156 µm long; Japan, China, Korea ...... P. composita Zahlbr. 8a. Norstictic or salazinic acid present ...... 9

9. Salazinic acid present ...... 10 9a. Norstictic acid present ...... 11

10. Spores 50-55 µm long; China ...... P. bambusetorum Zahlbr. 10a. Spores 80-100 µm long; Costa Rica ...... Lepra amnicola (Elix & A.W.Archer) I. Schmitt, Hodkinson & Lumbsch [Note: norstictic acid in Aus.] ...... [P. amnicola Elix & A.W.Archer]

11. Spores to 120-300 µm long; disc black; Eur...... P. rhodiensis Erichs. 11a. Spores 80-145 mm long ...... 12

12. Spores 80-100 µm long; Aus ...... Lepra amnicola (Elix & A.W.Archer) I. Schmitt, Hodkinson & Lumbsch ...... [P. amnicola Elix & A.W.Archer] 12a. Spores 110-145 µm long ...... 13

13. Spores 120-145 µm long; discs 0.8-1.3 mm diam.; Aus., Thailand ...... Lepra asiana (Vain.) A.W.Archer & Elix ...... [P. asiana Vain.] 13a. Spores 110-135 µm long; discs 0.5-0.75 mm diam.; S.Am...... P. purpurascens Müll.Arg.

14. Lecanoric or gyrophoric acid present ...... 15 14a. 2-O-Methylconfluentic, planaic or thamnolic acid present ...... 16

15. Lecanoric acid; spores 105-125 µm long; Philippines; PNG ...... P. philippina Vain. 15a. Gyrophoric acid present; spores 130-162 µm long; NZ ...... P. murrayi Elix & A.W.Archer

16. Thamnolic acid present ...... 17 16a. 2-O-Methylconfluentic or planaic acid present ...... 18

17. Spores 100-140 µm long; Japan ...... P. subcomposita Oshio 17a Spores to 94 µm long; N. Am...... Lepra trachythallina (Erichs.) Lendermer & R.C. Harris ...... [P. trachythallina Erichs] [Spores 130 µm long; OTA 61764; NZ; ...... Pertusaria sp.]

18. Planaic acid present; spores 100-180 µm long; China...... P. paraquilianensis Q. Ren & Z.T. Zhao 18a. 2-O-Methylconfluentic acid present; spores 50-140 µm long; Arctic alpine ...... P. saximontana Wetmore 

Group 10 Apothecia, present disciform; ascospores 2 per ascus; on rock

1. Thiophaninic and stictic acids present ...... 2 1a. Thiophaninic acid absent; depsidones or depsides present ...... 4

2. Spores with rough inner walls ...... P. pluripuncta Nyl. 2a. Spores with smooth inner walls ...... 3

3. Spores 60-80 µm long; on moss; N.Am ...... P. flavicunda Tuck. 3a. Spores 48-60 µm long; on moss; N.Am Arctic ...... P. atra Lynge

19 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

4. Depsidones [norstictic, protocetraric or salazinic acids] present ...... 5 4a. Depsides [lecanoric, 2'-O-methylperlatolic or thamnolic acids] present ...... 20

5. Protocetraric acid present ...... 6 5a. Norstictic or salazinic acid present ...... 7

6. Spores 100-126 µm long; disc white pruinose; thallus esorediate; Eur...... P. pseudoparotica Sipman 6a. Spores 100-150 µm long; thallus sorediate; Eur ...... P. digrediens Nyl.

7. Salazinic acid present; spores 120-150 µm long; Greece ...... P. parotica Sipman 7a. Norstictic acid present ...... 8

8. Norstictic acid only present ...... 9 8a. Norstictic and protocetraric or picrolichenic acids present ...... 14

9. Spores ≥110µm long ...... 10 9a. Spores ≤110 µm long ...... 12

10 Europe; spores 110-160 µm long...... P. pentelici J. Steiner 10a. South America ...... 11

11. Spores 120-175 µm long...... P. albidopallens Nyl. 11a. Spores 110-115 µm long...... P. scutellina Nyl

12. Europe; spores 80-100 µm long, becoming grey brown; Eur ...... P. lactescens Mudd 12a. North or South America ...... 13

13a. South America; spores 88-92 µm long ...... P. lecanorina Nyl. 13b. North America; spores 75-110 µm long ...... P. brattiae Lumbsch & T.H. Nash

14(8a).Norstictic and protocetraric acids present; spores 80-150 µm long; N.Afr ...... P. fdoulesiana Harm. 14a. Norstictic and picrolichenic acids present ...... 15

15. Spores < 85 µm long ...... 16 15a. Spores .> 85 µm long ...... 18

16. Spores 70-85 µm long [110-115 µm long; S.Afr...... Lepra leonina (Stizenb.) I. Schmitt, Hodkinson & Lumbsch ...... [P. leonina Stizenb.] 16a. Spores ≤ 70 µm long ...... 17

17. Spores 67-70 µm long; S.Afr...... P. wawreanoides Nyl. 17a. Spores 46-56 µm long; S.Afr., S.Am ...... P. subdealbata Nyl.

18 North Africa; spores 80-150 µm long ...... P. fdoulesiana Harm. 18a. South America ...... 19

19. Spores 162-250 µm long ...... Lepra rugifera (Müll.Arg.) I. Schmitt, Hodkinson & Lumbsch ...... [P. rugifera Müll.Arg.] 19a. Spores 84-125(-150) µm long ...... P. alterimosa Darb.

20(4a) Thamnolic acid present; spores 80-110 µm long; N.Am, China, Eur...... Lepra trachythallina (Erichs.) Lendemer & R.C. Harris ...... [P. trachythallina Erichs.] 20a. Lecanoric acid or 2'-O-methylperlatolic acid present ...... 21

21. Lecanoric acid present[?]C+red; spores 140-160 µm long; Asia P. haematommoides Zahlbr. 21a. 2'-O-Methylperlatolic acid present; also on soil, moss; spores 50-160 µm long; Arctic alpine

20

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

...... Pertusaria christae Dibben & Poelt

Group 11. Apothecia present, verruciform; ascospores 2 per ascus, with smooth walls; on rock

1. 4,5-Dichlorolichexanthone, thuringione or thiophaninic acid present ...... 2 1a. Chlorinated xanthones absent ...... 11

2. Thuringione or thiophaninic acid present ...... 3 2a. 4,5-Dichlorolichexanthone present ...... 4

3. Thuringione present; spores 110-150 µm long; Japan, Korea ...... P. astomoides Nyl. 3a. Thiophaninic and norstictic acids present; spores 175–250 x 70–100 µm; SW Afr...... P. salax Brusse

4. 4,5-Dichlorolichexanthone and 2'-O-methylperlatolic acid present; spores 63-135 µm long .. China, on alpine moss over rock ...... P. albiglobosa Q.Ren 4a. 4,5-Dichlorolichexanthone and stictic or norstictic acids present ...... 5

5. 4,5-Dichlorolichexanthone and norstictic acid present; spores 97-105 µm long; ...... ostioles black, conspicuous; West Indies ...... P. praetervisa var. expallens Vain 5a. 4,5-Dichlorolichexanthone and stictic acid present ...... 6

6. Thallus isidiate; spores 170-230 µm long; Eur ...... P. areolata (Ach.) A.Massal. 6a. Thallus not isidiate ...... 7

7. Spores 100-230 µm long ...... 8 7a. Spores 80-115 µm long ...... Spores 95-115 µm long; ostioles black; PNG ...... P. atrospilota A.W.Archer & Eli Spores 82-112 µm long; ostioles translucent; West Indies P. praetervisa var. pileolata Vain. Spores 92-102 µm long; ostioles black, punctiform; West Indies ...... P. praetervisa var. straminea Vain.

8. Spores 155-230 µm long ...... 9 8a. Spores 100-175 µm long ...... 10

9. Spores 155-230 µm long; ostioles black; Eur, Japan, Korea; also on wood ...... P. pertusa (L) Tuck. 9a. Spores 170-215 µm; medulla I+ blue; Corsica ...... P. subrupestris Zschacke

10. Spores 100-170 µm long; Russia ...... P. globulata Oxner & A.M.Volkova 10a. Spores 140-175 µm long; ostioles black; Aus, PNG ...... P. vulpina A.W.Archer

11. Thallus K-; 2'-O-methylperlatolic acid present ...... 12 11a. Thallus K+ red and C- or K+ red and C+ red ...... 13

12. Spores 130-150 µm long; thallus isidiate; Antarctic ...... P. signyae Øvstedal 12a. Spores 100-150 µm long; thallus non-isidiate; with confluentic and planaic acids present ..... India ...... P. indica Srivasrava & Awasthi ...... ?syn:P. kodaikanalensis Choisy

13. Thallus K+ red, C+ red; alectorialic acid present; spores 22-40 µm; Arctic alpine ...... P. geminipara (Th.Fr.) C.Knight ex Brodo 13. Thallus K+ red, C-ve; norstictic acid present ...... 14

14(12). Norstictic acid only present ...... 15 14a. Norstictic and protocetraric acids present; spores 80-150 µ long; N.Afr.P. fdoulesiana Harm.

15. Spores 100-140 µm long; S.Am...... P. tesselaria Müll.Arg. 15a. Spores 70-100 µm; Japan ...... P. pachyplaca Nyl.

21 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

Spores 110-150 µm; Japan ...... P. rhagadoplaca Nyl. Spores 120-250 µm long ...... 16

16. Thallus isidiate; spores 125-200 µm long; Eur ...... P. pseudocorallina (Lilj.) Arnold 16a. Thallus not isidiate...... 17

17. Thallus shiny; spores 145-270 µm long; US...... P. californica Dibben 17a. Thallus not shiny ...... 18

18 Spores 150-250 µm long; Eur...... P. apennina Bagl. . Spores 140-200 µm long; Hawaii ...... P. zahlbruckneri H. Magn. Spores 150-210 µm; India, with atranorin ...... Pertusaria sp. Singh & Sinha Spores120-200 µm long [85-245 µm, Dibben]; on moss; N.Am, Arctic ...... P. coriacea (Th.Fr.) Th.Fr.

Group 12. Apothecia present, verruciform; ascospores 2 per ascus, with rough walls; on rock

1. 4,5-Dichlorolichexanthone present ...... 2 1a. 4,5-Dichlorolichexanthone absent ...... 4

2. Divaricatic acid present; spores 120-192 µm long ...... P. flindersiana Kantvilas & Elix 2a. Norstictic or stictic acids present ...... 3

3. Norstictic acid present; spores 140-200 µm long; Aus, NZ ...... P. knightiana Müll.Arg. 3a. Stictic acid present; spores 150-180 µm long; Indonesia ...... P. tjibodana Zahlbr.

4. Norstictic acid absent; perlatolic and stenosporic acids present; spores 175-210 µm long; NZ [cf. corticolous P. sorodes Stirt]...... P. allanii Zahlbr. 4a. Norstictic acid present ...... 5

5. Norstictic and stenosporic acids present; spores 153-223 µm long; N.Am; China ...... P. plittiana Erichs. 5a. Norstictic acid only present ...... 6

6. Spores 190-245(-280) µm long; epithecium K-ve; NZ ...... P. subverrucosa Nyl. 6a. Spores 150-250 µm long; epithecium K+ violet; Eur...... P. appenina Bagl. ex Massal.

Group 13. Apothecia present, verruciform; ascospores 2 per ascus, with smooth walls; on bark or wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 46

2. Lichexanthone or derivatives present ...... 3 2a. Norlichexanthone derivatives or thiophaninic acid derivatives present ...... 7

3. Lichexanthone and stictic or norstictic acid ± depsides present ...... 4 3a. Chlorinated lichexanthones present ...... 28

4. Thallus isidiate ...... 5 4a. Thallus not isidiate; ...... 6

5. Spores 100-112 µm long, 2'-O-methylperlatolic and stictic acids present; Aus; PNG...... P. isidiosa A.W.Archer 5a. Spores 110-190 µm long; stictic acid present; Thailand ...... P. allomicrostoma Jariang.

22

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

6. Lichexanthone & norstictic acid present; spores 30-36 µm long [probably immature]; SAm ...... P. lichexanthonoimmersa Aptroot & M. Cáceres 6a. Lichexanthone & stictic acid present; spores 73-106 µm long; ostioles black; China ...... P. chinensis Müll.Arg.

7. Norlichexanthone derivatives [thiophanic acid, arthothelin, 6-O-methylarthothelin or ...... asemone] present ...... 8 7a. Thiophaninic acid and/or 2-chloro-6-O-methylnorlichexanthone present...... 9

8. Thiophanic acid, 6-O-methylarthothelin and stictic acid present; spores 90-100 µm long; LHI ...... P. malabara A.W.Archer & Elix 8a. Thiophanic acid, arthothelin and asemone present; spores 105-117 µm long; Aus...... P. saltuensis A.W.Archer & Elix

9. Thiophaninic acid present; 2-chloro-6-O-methylnorlichexanthone present or absent ...... 10 9a. Thiophaninic acid absent; 2-chloro-6-O-methylnorlichexanthone present ...... 27

10. Thiophaninic acid only or with 2'-O-methylperlatolic acid ...... 11 10a. Thiophaninic acid and depsidones [norstictic, stictic acids] present ...... 16 ......

11. Thiophaninic acid and 2'-O-methylperlatolic acid present; spores 87-110 µm long; S.Am...... P. cinerella Müll.Arg. 11a. Thiophaninic acid only present ...... 12

12. Canary Islands ...... 13 12a. New Zealand, Aus or SAm...... 14

13. Spores 100-180 µm long; also N.Afr, ...... P. tazzekensis Werner 13a Spores 125-182 µm long; ...... P. aceroae Hern.-Padr.et al.

14. Ostioles pale; spores 75-95 µm; Aus...... P. thiophaninica A.W.Archer 14a. Ostioles black; NZ or SAm ...... 15

15. Ostioles conspicuous; spores 87-102 µm long;; NZ ...... P. alboatra Zahlbr. 15a. Ostioles punctiform; spores 85 µm [Müll.Arg.]; SAm ...... P. carneola (Eschw.) Müll.Arg.

16(10a).Thiophaninic and norstictic acids present ...... 17 16a. Thiophaninic and stictic acids present ...... 19

17. Thiophaninic and norstictic acids ±depsides present; spores 70-110 µm; Aus...... [also 3-4 per ascus] ...... P. trimera (Müll.Arg.) A.W.Archer 17a. Thiophaninic and norstictic acids only present ...... 18

18. Spores 70-95 µm long; Spain ...... P. luteola Boqueras 18a. Spores 86-118 µm long; N.Am ...... P. neolecanina Lumbsch & Nash

19. Spores 80-105 µm long; Aus, Galapagos, Korea ...... P. thiospoda C. Knight 19a. Spores otherwise ...... 20

20. Spores 70-110 µm long; Japan ...... P. subrugosa Nyl. 20a. Spores >110 µm long ...... 21

21. Spores 100-187 µm long ...... 22 21a. Spores <100 µm or hypostictic acid present ...... 24

22. Eur, Macaronesia; spores 140-145 µm long; ...... P. ficorum Zahlbr. 22a. Socotra ...... 23

23 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

23. Spores 100-145 µm long; ...... P. cicatricosa Müll.Arg. 23a. Spores 145-187 µm long; ...... P. schizostoma Müll.Arg.

24. Thiophaninic and stictic acids; ...... 25 24a. Thiophaninic and stictic acids with additional hypostictic acid present ...... 26

25. Spores 75-82 µm long; S.Am...... P. montevidensis Zahlbr. 25a. Spores 125-150 µm long; Aus...... P. epacrospora A.W.Archer

26. Spores 87-125 µm long; S.Afr...... P. dispersa Vain.  26a. Spores 60-90 µm long; West Indies ...... P. antillarum Vain.

27. 2-Chloro-6-O-methylnorlichexanthone and 2'-O-methylperlatolic acid present; ...... spores 80-100 µm; S.Am...... P. cinerella var. pampeana A.W.Archer & Messuti 27a. 2-Chloro-6-O-methylnorlichexanthone and stictic acid present; spores 87-100 µm long; ...... Eur. N.Am, S.Afr, Asia, Aus ...... P. pustulata (Ach.) Duby

28(3a) 2,4- & 2,5-Dichlorolichexanthones and 2,4,5-trichlorolichexanthone present ...... 29 28a. 4,5-Dichlorolichexanthone present ...... 34

29. Polychlorolichexanthones only present; spores 110-130 µm; Aus...... P. xanthonaria A.W.Archer & Elix 29a. Polychlorolichexanthones with 2’-O-methylperlatolic acid and/or stictic acid ...... 30

30. 2’-O-Methylperlatolic acid present; spores 75-160 µm; Japan ...... P. denotanda Nyl. 30a. Stictic acid present...... 31

31. Stictic acid and 2’-O-methylperlatolic acid present; spores 80-120 µm long; PNG ...... P. damiensis A.W.Archer & Elix 31a. Stictic acid present, 2’-O-methylperlatolic acid absent ...... 32

32. Thallus sorediate; spores 140-168 µm long; Thailand .... P. pertusella var. sorediata Jariang. 32a. Thallus esorediate ...... 33

33. Spores 100-150 µm long; pantropical? ...... P. pertusella Müll.Arg. 33a. Spores 90-110 µm; Cook Islands ...... P. homilocarpa A.W.Acher & Elix

34(28a). 4,5-Dichlorolichexanthone only present; spores 120-155 µm long; Aus...... P. irregularis Müll.Arg. 34a. 4,5-Dichlorolichexanthone, norstictic acid, stictic acid ± depsides, or depsides present ...... 35

35. 4,5-Dichlorolichexanthone and norstictic acid present; spores 86-122 µm long; Thailand ...... P. neoknightiana Jariang. [cf. saxicolous P. praetervisa var. expallescens Vain. West Indies] [with additional 2-O-methylstenosporic and 2-O-methylperlatolic acids; ...... spores 130-150 µm long; 53470 (M) ...... P. "chiodectonoides"] 35a. 4,5-Dichlorolichexanthone, depsides or stictic acid ± depsides present ...... 36

36. 4,5-Dichlorolichexanthone and depsides present ...... 37 36a. 4,5-Dichlorolichexanthone and stictic acid ± depsides present ...... 41

37. 4,5-Dichlorolichexanthone and 2'-O-methylstenosporic acid; spores 100-110 µm long; Aus...... P. stenospora A.W.Archer & Elix

37a. 4,5-Dichlorolichexanthone and 2'-O-methylperlatolic acid or 2-O-methylperlatolic acid present ...... 38

38. 4,5-Dichlorolichexanthone and 2-O-methylperlatolic acid present ...... 39 38a. 4,5-Dichlorolichexanthone and 2'-O-methylperlatolic acid present ...... 40

24

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

39. Spores 105-135 µm long; ostioles translucent; Aus...... P. meeana A.W.Archer & Elix 39a. Spores 105-137 µm long; ostioles black; Thailand ...... P. parameeana Jariang.

40. Spores 110-135 µm long; Aus, PNG, New Cal., China ...... P. pycnothelia Nyl. 40a. Spores 65-95 µm long; Israel; ...... P. carmelii Reichert & Galun

41(36a).4,5-Dichlorolichexanthone, stictic acid and 2'-O-methylperlatolic acids present ...... 42 41a. 4,5-Dichlorolichexanthone and stictic acid present ...... 43

42. Spores 137-160 µm long; Aus, Fiji ...... Pertusaria queenslandica Elix & A.W.Archer 42a. Spores 65-80 µm long; Philippines ...... P. striolata Räsänen

43. Spores 145-230 µm long; Eur. ?widespread; also on rock ...... P. pertusa (Weigel) Tuck. 43a. Spores shorter ...... 44

44. North America; spores 120-180 µm long; ...... P. consocians Dibben 44a. Australia ...... 45

45. Ostioles black; spores 82-112 µm long; Aus, PNG, Sri LankaP. pseudococcodes Müll.Arg. 45a. Ostioles pale; spores120-150 µm long; Aus...... P. quadraginta A.W.Archer & Elix

46(1a) Lichen compounds absent ...... 47 46a. Lichen compounds present ...... 51

47. Spores <100 µm long ...... 48 47a. Spores >100µm long ...... 49

48. Spores 64-100 µm long; S.Am...... P. albidella Nyl. 48a. Spores 50-60 µm long; S.Am...... P. cryptocarpoides Vain.

49. Spores 210-250 µm long; NZ ...... P. hadrospora A.W.Archer & Elix 49a. Spores < 200µm long ...... 50

50. Spores 110-125 µm long; Aus, NZ...... P. melaleucoides Müll.Arg. 50a. Spores 60-120 µm long; circumboreal ...... P. stenhammerii Hellb.

51(46a). Depsides and/or depside methyl esters present ...... 52 51a. Depsidones [fumarprotocetraric, norstictic, protocetraric, stictic acids] present ...... 56

52. Depside methyl esters present ...... 53 52a. Depside methyl esters absent; depsides present ...... 54

53. Methyl 2'-O-methylstenosporate present; spores 140-174 µm; Thailand ...... P. archeri Jariang. 53a. Divaricatic acid and methyl 2,2'-di-O-methyldivaricatate present; spores 90-105 µm long; ... Taiwan ...... P. platycarpiza Zahlbr.

54. Planaic acid present; spores 90-130 µm long; China. P. paraquilianensis Q.Ren & Z.T.Zhao 54a. Divaricatic acid derivatives or 2'-O-methylhyperphyllinic acid present ...... 55

55. 2'-O-Methylstenosporic, 2'-O-methyldivaricatic and 2,2'-di-O-methyldivaricatic acids present; spores 165-215 µm long; S.Am ...... P. velloziae (Vain.) Erichs. 55a. 2'-O-Methylhyperphyllinic acid present; spores 100-135 µm long; Japan ... P. yasudae Vain.

56(51a).Protocetraric or fumarprotocetraric acid present; ...... 57 56a Norstictic or stictic acid present ...... 59

57. Fumarprotocetraric acid present; spores 105-120 µm long; Japan ...... P. subpustulata Nyl. 57. Protocetraric acid present ...... 58

25 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

58. Spores 100-120 µm long; Aus...... P. aphelospora (A.W.Archer) A.W.Archer & Elix 58a. Spores 40-70 µm long [fide Nyl.]; Japan ...... P. diffidens Nyl.

59(56a). Stictic acid present ...... 60 59a. Norstictic acid present ...... 61

60. Spores 80-120 µm long; Aus, Mexico ...... P. porinella Nyl. 60a. Stictic and 2'-O-methylperlatolic acids present; spores 100-170 µm long; Thailand ...... P. subcopelandii Jariang.

61. Norstictic acid only present ...... 62 61a. Norstictic acid and depsides present ...... 66

62. Spores 200-360 µm long; Asia ...... P. oshioi Wei 62a. Spores < 200 µm long ...... 63

63. Asia ...... 64 63a. Europe or South America ...... 65

64. India; spores 70-157µm long; [+atranorin] ...... P. hirtella Singh & Sinha 64a. Japan; spores 130-145 µm long; ...... P. mendax Müll.Arg.

65. Europe; spores 110-195 µm long; ...... P. servitiana Erichs. 65a. South America; spores 110-140 µm long; K+ red (fide Vainio 1900) ...... P. pycnothelioides Vain.

66(61a). Norstictic and 2-O-methylstenosporic or divaricatic acids present ...... 67 66a. Norstictic and perlatolic, ±stenosporic acids present...... 68

67 Norstictic and 2-O-methylstenosporic present; spores 110-137 µm long; India ...... P. wattiana Müll.Arg. 67a. Norstictic and divaricatic acids present; spores 75-100 µm long; China ...... P. subrosacea Zahlbr.

68. Spores 120-170 µm long; Aus ...... P. hartmannii Müll.Arg. 68a. Spores 120-125 µm long; Japan, China, Korea; rarely ± perlatolic acid ...... cf. P. mendax, above ...... P. subobductans Nyl.

Group 14. Apothecia present, verruciform; ascospores 2 per ascus, rough; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 13

2. Norlichexanthone derivatives present ...... 3 2a. Lichexanthone and derivatives present ...... 6

3. Arthothelin and 6-O-methylarthothelin present; spores 80-110 µm long; Eur...... P. dispar J. Steiner 3a. Thiophaninic and stictic acids present ...... 4

4. Socotr; spores 140-165 µm long; ...... P. subflavens Müll.Arg. 4a. Northern Hemisphere ...... 5

5. Spores 70-150 µm long; N.Am, China, Korea ...... P. xanthodes Müll.Arg. 5a. Spores 90-120 µm long; Eur...... P. heterochroa (Müll.Arg.) Erichs.

6. Lichexanthone present; spores 170-225 µm long; S.Am.P. tetrathalamia var. major Müll.Arg. [also 1 per ascus]

26

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

6a. Chlorolichexanthones present ...... 7

7. 4-Chlorolichexanthone and 2-O-methylperlatolic acid present; spores 190-250 µm long; NZ ...... P."communis" sens. Knight (M) 7a. Polychlorolichexanthones present ...... 8

8. 4,5-Dichlorolichexanthone present ...... 9 8a.. Other polychlorolichexanthones present ...... 10

9. 4,5-Dichlorolichexanthone and perlatolic and stenosporic acids present; ...... spores 160-240 µm long; NZ ...... P. sorodes Stirt. [cf. saxicolous P. allani Zahlbr.] ...... 9a. 4,5-Dichlorolichexanthone and 2'-O-methylperlatolic acid present; spores 95-120 µm long. Aus; PNG ...... P. trachyspora A.W.Archer

10. 2,4- & 2,5-Dichlorolichexanthone, 2,4,5-trichlorolichexanthone and stictic acid present ... 11 10a. 2,5-Dichlorolichexanthone, 2,4,5-trichlorolichexanthone, 2'-O-methylperlatolic and ...... 2-O-methylperlatolic acids present; spores 140-175 µm long; Aus...... P. elliptica Müll.Arg. var. bispora Elix & A.W.Archer 11. Thallus lacking isidia & soredia; spores 100-150 µm long; ?circumtropical ...... P. cicatricosa Müll.Arg. 11a. Thallus with isidia or soralia ...... 12

12. Thallus isidiate; spores 116-180 µm long; Thailand ...... P. takensis Jariang. & A.W.Archer 12a. Thallus sorediate; spores 126-204 µm long; Thailand ...... P. langsangensis Jariang. & A.W.Archer

13(1a) Depsidones and ±depsides present ...... 14 13a. Depsidones absent; depsides present ...... 25

14. Depsidones [stictic or norstictic acid] and depsides present ...... 15 14a. Depsidones only present ...... 19

15. Stictic and 2'-O-methylperlatolic acids present; spores 150-175µm long; PNG ...... P. bundiensis A.W.Archer & Elix 15a. Norstictic acid present ...... 16

16. India ...... 17 16a. Asia or North America ...... 18

17. Norstictic and divaricatic acids present; spores 120-166 µm long; PNG, Thailand ...... P. allothwaitesii Jariangpraseret & A.W.Archer 17a. Norstictic ±planaic acids present; spores 100-160 µm long; N.Am ...... P. neoscotia Lamb

18. Norstictic and 2-O-methylstenosporic acids present; spores 120-137 µm long, rough; India ...... P. wattiana Müll.Arg. 18a. Norstictic acid and atranorin present; spores 70-157 µm long; India ...... P. hirtella Singh & Sinha

19. Hypoprotocetraric, fumarprotocetraric or protocetraric acids present ...... 20 19a. Stictic, norstictic acid or variolaric acids present ...... 22

20. Hypoprotocetraric acid present; spores 125-175 µm long; NZ .. P. vallicola Elix & Malcolm 20a. Fumarprotocetraric or protocetraric acids present ...... 21

21. Fumarprotocetraric acid present; spores 75-135 µm long; N.Am, China P. subpertusa Brodo 21a. Protocetraric acid present; spores 110-160 µm long; Aus, PNG, Korea, Asia ...... P. thwaitesii Müll.Arg.

22. Variolaric acid present; spores 100-120 µm long; N.Am ...... P. obruta Harris

27 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

22a. Stictic or norstictic acids present ...... 23

23. Stictic acid present...... 24 23a. Norstictic acid present; spores 150-200 µm long; PNG . P. perthwaitesii A.W.Archer & Elix

24. Stictic acid present; spores 110-125 µm long; PNG ...... P. cicatricosa Müll.Arg. var. deficiens A..W. Archer & Elix 24a. Stictic and menegazziaic acids present; spores 150-175 µm long; India ...... P. neilgherrensis (Müll.Arg.) Awasthi & Srivastava

25(13a).Methyl 2-O-methyldivaricatate and 2-O-methyldivaricatic acid present; ...... spores 86-150 µm long; Thailand ...... P. litchicola Jariang. & A.W.Archer 25a. 2'-O-Methylperlatolic acid present; spores 185-250 µm long; PNG ...... P. subsorodes Elix & A.W.Archer

Group 15. Apothecia present, verruciform; ascospores 4 per ascus, with smooth walls; on rock or moss

1. Lichexanthone and stictic acid present; spores70-85 µm long; Central Afr...... P. rhodesica Vain. 1a. Lichexanthone absent ...... 2

2. Chlorinated xanthones present ...... 3 2a. Chlorinated xanthones absent ...... 14

3. Thiophaninic acid present ...... 4 3a. Chlorinated lichexanthones (such as 4,5-dichlorolichexanthone) present ...... 6

4. Thiophaninic, confluentic and planaic acids present; spores 100-130 µm long; NAm ...... P. tejocotensis de Lesd. 4a. Thiophaninic and stictic acids present; Japan ...... 5

5. Anthraquinone absent; spores 100-130 µm long ...... P. boninensis Shibuchi 5a. Anthraquinone present; spores 100-160 µm long ...... P. kashiwadanii Shibuchi

6. 2-Chlorolichexanthone, stictic and 2-O-methylsuperlatolic acids present; ...... spores 108-125 µm long; PNG ...... P. aptrootii A.W.Archer & Elix

6a. 4,5-Dichlorolichexanthone or 4-chlorolichexanthone present ...... 7

7. 4,5-Dichlorolichexanthone present ...... 8 7a. 4-Chlorolichexanthone present; spores 100-120 µm long; Antarctic, Marion Is...... [? not published; not in Index Fungorum 21.2.2018] ...... "P. gremmenii Øvstd."

8. Growing on moss ...... 9 8a. Growing on rock ...... 12

9. 4,5-Dichlorolichexanthone, stictic and 2-O-methylperlatolic acids present; ...... spores 140-175 µm long; S.Am...... P. tapadensis Elix & A.W.Archer 9a. 4,5-Dichlorolichexanthone and stictic acid present ...... 10

10. Spores (1-)2(-3)/ascus, 143 µm long; also on wood; N.Am, arctic ...... P. subobducens Nyl. 10a. Spores <143 µm long ...... 11

11. Spores 93-104 µm long; Japan, China, Korea ...... P. quartans Nyl. 11a. Spores 42-96 µm long; Arctic ...... P. trochisea Norm.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

12(8a) 4,5-Dichlorolichexanthone only present; spores 2-4 per ascus, 70-90 µm long; Cape Verde Is ...... P. aleianta Nyl. 12a. 4,5-Dichlorolichexanthone with stictic or norstictic acids ...... 13

13. 4,5-Dichlorolichexanthone and norstictic acid present; spores 80-90 µm long; S.Am ...... P. rudecta Müll. Arg. 13a. 4,5-Dichlorolichexanthone and stictic acid present; spores 80-105 µm long; Aus...... P. trevethensis A.W.Archer

14(2a).On moss; norstictic acid present ...... 15 14a. On rock ...... 16

15. On moss; spores 50-180 µm long; circumpolar...... P. glomerata (Ach.) Schaer. 15a. On moss and rock; [also 2,3 or 5/ascus], 80-230 µm long; polar ...... P. alaskensis Erichs.

16. 2-O-Methylperlatolic or confluentic acid present ...... 17 16a. Norstictic acid present ...... 18

17. 2-O-Methylperlatolic acid present; spores 145-185 µm long; S.Am...... P. spegazzinii Müll.Arg. 17a. Confluentic acid present; spores 108-130 µm long; S.Am ...... P. malvinae Messuti & A.W.Archer

18. Spores 55-75 µm long; Mauritius ...... P. hymenelioides J.C.David 18a. Spores 60-90 µm long; NZ ...... P. graphica C.Knight

Group 16. Apothecia present, verruciform; ascospores 4 per ascus, rough; on rock So far, no taxa are known in this Group.

Group 17. Apothecia present, verruciform; ascospores 4 per ascus, smooth; on wood

1. Xanthones present ...... 2 1a. Xanthones absent ...... 52

2. Lichexanthone or derivatives present ...... 3 2a. Norlichexanthone derivatives present [including thiophaninic acid] ...... 41

3. Lichexanthone present ...... 4 3a. Chlorinated lichexanthone derivatives present ...... 11

4. Lichexanthone only present; spores 80-120 µm long; N.Am, Central Afr...... P. valliculata Dibben 4a. Lichexanthone and stictic acid ± depsides or 2-chlorolichexanthone present ...... 5

5. Lichexanthone and 2-chlorolichexanthone present ...... 6 5a. Lichexanthone and stictic acid ± depsides present ...... 7

6. With additional 2-O-methylsuperlatolic and 2-O-methylhyperlatolic acids; spores 90-115 µm long; Aus, S.Am, Cuba, PNG ...... P. depressa (Fée) Müll.Arg. 6a, With additional stictic acid; spores 85-120 µm long; Aus...... P. atromaculata A.W.Archer & Elix

7. Lichexanthone, constictic and stictic acids present; spores 75-130 µm long; S.Am ...... P. rhodiza Nyl. 7a. Lichexanthone, stictic acid and depsides present ...... 8

8. Lichexanthone, stictic and confluentic acids present...... 9

29 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

8a. Lichexanthone, stictic acid and stenosporic acid derivatives present ...... 10

9. Spores 2-4/ascus, 80-176 µm long; Thailand ...... P. inthanonensis Jariang. 9a. Spores 4-6-8/ascus, 87-110 µm long; S.Am...... P. gracilis var. heteromera Müll.Arg.

10. 2'-O-Methylstenosporic acid present; spores 90-110 µm long; Thailand ...... P. elixii Jariang. 10a. 2,2'-Di-O-methylstenosporic acid present; spores (3-)4/ascus 76-92 µm long; Thailand ...... P. alboaspera var. tetraspora Jariang.

11(3a). 2-Chlorolichexanthone, stictic, 2-O-methylisohyperlatolic and 2-O-methylsuperlatolic acids present; spores 105-125 µm long; Aus...... P. follmanniana A.W.Archer & Elix 11a. 4.5-Dichlorolichexanthone or 2,4,5-trichlorolichexanthone present ...... 12

12. 4,5-Dichlorolichexanthone present ...... 13 12a. 2,4,5-Trichlorolichexanthone present ...... 38

13. 4,5-Dichlorolichexanthone only present; ...... 14 13a. 4,5-Dichlorolichexanthone and depsidones and/or depsides present ...... 15

14. Spores 75-110 µm long; ostioles inconspicuous, pale brown, translucent; S.Am ...... P. platystoma Malme 14a. Spores 75-100 µm long; ostioles inconspicuous, black; S.Am ...... P. simulans Malme

15. 4,5-Dichlorolichexanthone and norstictic acid present; spores 80-110 µm long; S.Am; ...... [H-NYL 3702 pm] ...... P. "rhodiza" Nyl. 14a. 4,5-Dichlorolichexanthone and stictic acid or depsides present ...... 15

15. 4,5-Dichlorolichexanthone and depsides or constictic acid present ...... 16 15a. 4,5-Dichlorolichexanthone and stictic acid ± depsides present ...... 25

16. 4,5-Dichlorolichexanthone and depsides present ...... 17 16a. 4,5-Dichlorolichexanthone and constictic acid present; spores 100-125 µm ; Brazil ...... P. marcellii A.W.Archer & Elix

17. 4,5-Dichlorolichexanthone and stenosporic acid derivatives present ...... 18 17a. 4,5-Dichlorolichexanthone and perlatolic acid derivatives present ...... 19

18. 4,5-Dichlorolichexanthone and 2,2'-di-O-methylstenosporic acid present; spores 2-3(-4)/ascus 84-142 µm long; Thailand ...... P. subplanaica var. tetraspora Jariang. & A.W.Archer 18a. 4,5-Dichlorolichexanthone and 2'-O-methylstenosporic acid; spores 80-100 µm; Thailand ...... P. kansrii Jariang.

19. Species present in Thailand ...... 20 19a. Species present elsewhere ...... 22

20. 4,5-Dichlorolichexanthone and planaic acid or derivatives present ...... 21 20a. 4,5-Dichlorolichexanthone and 2'-O-methylanziaic acid present; spores 84-146 µm long; ...... P. phusoidaoensis Jariang..

21. 4,5-Dichlorolichexanthone and planaic acid present; spores (1-)2-3/ascus 90-174 µm long; Thailand ...... P. siamensis Jariang. 21a. 4,5-Dichlorolichexanthone and 4-O-demethylplanaic acid present; spores 100-156 µm long; ...... P. khuntanensis Jariang.

22. Species present in Australia...... 23 22a. Species present in Papua New Guinea or Vanuatu ...... 24

23. 4,5-Dichlorolichexanthone, planaic and 2-O-methylperlatolic acids present; spores 4/ascus,

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

95-125 µm long; Aus ...... P. doradorensis Elix & A.W.Archer 23a. 4,5-Dichlorolichexanthone and 2-O-methylperlatolic acid present; spores 80-94 µm long; Aus, S.Am ...... P. malmeii Elix & A.W.Archer

24a. Vanuatu; 4,5-dichlorolichexanthone and 2-O-methylperlatolic acid present; ...... spores 64-78 µm long; Vanuatu ...... P. wilsoniana A.W.Archer & Elix 24. Papua New Guinea; 4,5-dichlorolichexanthone and 2-O-methylisohyperlatolic present; spores 105-115 µm; PNG ...... P. streimannii Elix & A.W.Archer

25. 4,5-Dichlorolichexanthone and stictic acid only present ...... 26 25a 4,5-Dichlorolichexanthone, stictic acid and depsides present ...... 31

26. Thallus isidiate; spores 90-160 µm long; Eur...... P. coronata (Ach.) Th.Fr. 26a, Thallus not isidiate ...... 20

27 Spores >100 µm long ...... 28 27a. Spores <100 µm long ...... 29

28. Spores 150-185 µm long, (2-)4(-5) per ascus; Japan ...... P. glauca Zahlbr. 28a. Spores 112-142 µm long; Fiji ...... P. athrocarpar var. deficiens.Elix & A.W.Archer

29. Spores 2-3(-4)/ascus; 75-100 µm long; Tahiti; verrucae confluent ...... P. hypochrysea Vain. 29a. Species in the Northern Hemisphere ...... 30

30. Spores 2-4/ascus, 51-140 µm long; pantemperate, N. hemisphere; Hawaii ...... verrucae flattened ...... P. leucostoma (Bernh.) Massal. 30a.. Spores 60-90 µm; verrucae contricted at base; Eur ...... P. caesioumbrina Eitner

31. 4,5-Dichlorolichexanthone, stictic acid and perlatolic or stenosporic acid derivatives present ...... 33 31a. 4,5-Dichlorolichexanthone, stictic acid and confluentic acid or 2-O-methylconfluentic acid present; ...... 32

32. 4,5-Dichlorolichexanthone, stictic acid and 2-O-methylconfluentic acid present; ...... spores (3-)4/ascus, 85-125 µm long; PNG ...... P. bogia A.W.Archer & Elix ...... 32a. 4,5-Dichlorolichexanthone, stictic acid and confluentic acid present; spores 75-90 µm long; India ...... P. cinchonae Müll.Arg.

33. 4,5-Dichlorolichexanthone, stictic acid and perlatolic acid derivatives present ...... 23 33a. 4,5-Dichlorolichexanthone, stictic acid and 2'-O-methylstenosporic acid present; ...... spores 3-4/ascus, 80-108 µm long ...... P. thailandica Jariang.

34. 4,5-Dichlorolichexanthone, stictic acid and planaic acid present; spores 82-100 µm long; S.Am...... P. bonariensis Malme 34a. 4,5-Dichlorolichexanthone, stictic acid and 2-O-methylperlatolic or 2'-O-methylperlatolic acid present ...... 35

35. 4,5-Dichlorolichexanthone, stictic acid and 2'-O-methylperlatolic acid pesent ...... 36 35a. 4,5-Dichlorolichexanthone, stictic acid and 2-O-methylperlatolic acid present ...... 37

36. Spores 90-150 µm long; ostioles inconspicuous; Aus ...... P. hermaka A.W.Archer 36a. Spores 130-152 µm; ostioles black, conspicuous; Fiji ...... P. athrocarpa Elix & A.W.Archer

37. Spores (2-)4/ascus, 163-200 µm long; PNG, Philippines ...... P. copelandii Vain. 37a. Spores 4/ascus, 100-120 µm long; PNG ...... P. laeana A.W.Archer & Elix

38(12a) 2,4,5-Trichlorolichexanthone, 2-O-methylperlatolic and stictic acids; spores (2-)3(-4)/ascus, 70-100 µm long; Aus ...... P. tjaetabensis A.W.Archer & Elix 38a. 2,4,5-Trichlorolichexanthone and 2,5-dichlorolichexanthone present ...... 39

31 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

39. 2,4,5-Trichlorolichexanthone, 2,5-dichlorolichexanthone & 2-O-methylperlatolic acid present; spores 70-80 µm; Aus ...... P. aquilonia A.W.Archer & Elix 39a. 2,4,5-Trichlorolichexanthone, 2,5-dichlorolichexanthone and stictic acid present ...... 40

40. 2,4,5-Trichlorolichexanthone, 2,5-dichlorolichexanthone, stictic & confluentic acids present; spores (3-)4/ascus, 64-75 µm long; Aus ...... P. ewersii A.W.Archer & Elix 40a. 2,4,5-Trichlorolichexanthone, 2,5-dichlorolichexanthone and stictic acid present spores 95-125 µm long; Sri Lanka, Java, PNG, Aus...... P. ceylonica Müll.Arg.

41(2a) Arthothelin and derivatives present ...... 42 41a. Thiophaninic acid present ...... 43

42. 6-O-Methylarthothelin, arthothelin and stictic acid present; spores 117-125 µm long ...... circumtropical ? ...... P. quasiae (Fée) Nyl. 42a. 6-O-Methylarthothelin and arthothelin present; spores 45-62 µm long; PNG ...... P. inconspicua A.W.Archer & Elix

43. Thiophaninic acid ± stictic or norstictic acid or despsides ...... 44 43a. Thiophaninic acid, stictic acid and 4-O-methylisocryptochloropheaic acid or ...... anthraquinone pigment present ...... 51

44. Thiophaninic and norstictic acids present ...... 45 44a. Thiophaninic acid ± stictic acid or depsides ...... 46

45. Spores 68-110 µm long; N.Am ...... P. wulfenoides de Lesd. 45a. Spores 85-120 µm long; Socotra ...... P. socotrana Müll.Arg.

46. Thiophaninic acid and stictic acid present ...... 47 46a. Thiophaninic acid ± depsides ...... 48

47. Spores (3-)4/ascus, 75-90 µm long; Aus, China ...... P. abberans Müll.Arg. 47a. Spores 3(-4)/ascus,100-20 µm long; Aus...... P. ternata A.W.Archer & Elix

48. Thiophaninic acid only present; spores 4/ascus, 70-80 µm; New Caledonia, Thailand ...... P. endochroma Müll.Arg. 48a. Thiophaninic acid and depsides present ...... 49

49. Thiophaninic acid and 2'-O-methylperlatolic acid present; spores 4/ascus, 95-120 µm long: PNG ...... P. rechingeri Zahlbr. 49a. Species present in Australia...... 50

50. Thiophaninic acid and glomelliferic acid present; spores (2-)3/ascus, 80-120 µm long; Aus ...... P. glomellerifica Elix & A.W.Archer

50a. Thiophaninic acid with 2-O-methylperlatolic acid or 2,2'-di-O-methylstenosporic acid or confluentic acid present; spores 70-110 µm long; ostioles black, conspicuous; Aus ...... P. trimera (Müll.Arg.) A.W.Archer

51(43a).Unknown anthraquinone present [K+violet]; spores 4/ascus, 100-160 µm long; Japan ...... also saxicolous ...... P. kashiwadanii Shibuichi 51a. 4-O-Methylisocryptochloropheaic acid present; spores 4/ascus, 95-125(-145) µm long ...... P. paradoxica A.W.Archer & Elix var. tetraspora A.W.Archer & Elix

52(1a)Stictic or norstictic acid present ...... 53 52a. Stictic and norstictic acid absent ...... 60

53. Stictic acid or stictic acid with depsides present ...... 54 53a. Norstictic acid present ...... 58

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

54. Stictic acid present...... 55 54a. Stictic acid and stenosporic or perlatolic acid derivatives ...... 56

55. Spores 75-82 µm long, Qld...... P. dayi A.W.Archer & Elix 55a. Spores 140-160 µm; West Indies ...... P. dussii Vain..

56. Methyl 2'-O-methylstenosporate present; spores 2-3/ascus 118-150 µm long; Thailand ...... P. methylstenosporica Jariang. 56a. Stictic acid and perlatolic acid derivatives present ...... 57

57. 2-O-Methylperlatolic acid present; spores 2-(3-4)/ascus, 84-130 µm long; Thailand; Aus ...... P. loeiensis Jariang. 57a. 2'-O-Methylperlatolic acid present; spores (3-)4/ascus, 115-150 µm long; PNG ...... P. novaeguineae A.W.Archer & Elix

58(53a).Spores 85-110 µm long; Hawaii [K++red] ...... P. ruboreagens H. Magn. 58a. Spores 62-85 µm long ...... 59

59. Spores 62-85 µm; ostioles pale; S.Am, ...... P. cryptocarpa Nyl. 59a. Spores 65-85 µm; ostioles black; PNG ...... P. naduriensis A.W.Archer & Elix

60(52a).Lichen compounds present ...... 61 60a. Lichen compounds absent ...... 68

61. Atranorin present; spores 88-104 µm long; Thailand ...... P. phukaensis Jariang. 61a. Other depsides or protocetraric acid present ...... 62

62. 2-O-Methylsuperlatolic, 2'-O-methylisohyperlatolic or protocetraric acid present ...... 63 45a. Confluentic or 2'-O-methylperlatolic acid present ...... 66

63. Protocetraric acid present; spores 16-22 µm long; S.Am ...... P. huanicola Messuti & A.W.Archer 63a. Perlatolic acid or its derivatives present ...... 64

64. Perlatolic acid and fumarprotocetraric acid present; spores 90-150 µm long; Japan ...... P. nakamurae (Räs.) Dibben  64a. Derivatives of perlatolic acid present ...... 65

65. 2-O-Methylsuperlatolic acid present; spores 3-(4)/ascus, 80-100 µm long; PNG ...... P. manamensis A.W.Archer & Elix 65a. 2'-O-methylisohyperlatolic acid present; spores 65-69 µm long; Mexico ...... P. polystictoides Vain.

66. Confluentic acid present; spores 2-3(-4)/ascus, 72-134 µm long; Thailand ...... P. orientalis Jariang. 66a. 2'-O-Methylperlatolic acid present; ...... 67

67. Spores (2-)3/ascus, 140-168 µm long; China ...... P. parapycnothelia Q. Ren & Z.T. Zhao 67a. Spores 90-120 µm long; Cook Islands ...... P. atroguttata A.W.Archer & Elix

68(60a).In S.Am; spores 80-100 µm long; ...... P. polita var. tetramera Müll.Arg. 68a. In Australasia ...... 69

69. In PNG; spores 90-110 µm long; ostioles inconspicuous, translucent ...... P. plethocarpa A.W.Archer, Elix & Streimann 69a. In Norfolk Island; spores 95-115 µm long; ostioles black, lacking a hyaline margin ...... P. nebulosa A.W.Archer

Group 18. Apothecia present, verruciform; ascospores 4 per ascus, rough; on wood

33 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

1. Xanthones absent ...... 2 1a, Xanthones present ...... 8

2. Norstictic acid present; ...... 3 2a. Atranorin or stictic acid present ...... 4

3. Spores 84-144 µm long; S.Am...... P. reagens (Zahlbr.) Redinger 3a. Spores 90-120 µm long; ostioles black; West Indies...... P. plana Vain.

4. Atranorin only present; spores 88-104 µm long; Thailand ...... P. phukaensis Jariang. 4a. Stictic acid present...... 5

5. Stictic acid only present; ...... 6 5a. Stictic and 2'-O-methylperlatolic acid present; spores 70-80 x 36-40 µm; Qld...... P. gadgarrensis A.W.Archer & Elix

6. Spores 87-137(-165) µm long; ostioles punctiform; Java ...... P. javanica Müll.Arg. 6a. Spores 100-160 µm long ...... 7

7. Spores 140-160 µm long; ostioles becoming disciform; West Indies ...... P. dussii Vain. 7a. Spores 100-150 µm long; Japan ...... P. radiata Oshio

8. Lichexanthone present ...... 9 8a. Chlorinated lichexanthones present ...... 15

9. Lichexanthone and stictic or constictic acids present ...... 10 9a. Fatty acids present or lichexanthone alone ...... 14

10 Lichexanthone and stictic acid present ...... 11 10a. Lichexanthone and constictic acid present; spores [sl. rough] 125-160 µm long; S.Am...... P. ochrostoma Müll.Arg.

11. Lichexanthone and stictic acid present ...... 12 11a Lichexanthone, stictic acid and confluentic acid or 2-O-methyl- and ...... 2,4-di-O-methylolivetoric acids present ...... 13

12. Spores 97-125 µm long; S.Am...... P. tenella Müll.Arg. 12a. Spores 135-165 µm long; S.Am ...... P. tetrathalamia var. plicatula Müll.Arg.

13. With additional confluentic acid; spores 70-90 µm long; Tahiti .... P. trypetheliformis Nyl. 13a. With additional 2-O-methyl- and 2,4-di-O-methylolivetoric acid; spores 115-138 µm; ...... S.Am ...... P. ochrotheliza Nyl.

14. Lichexanthone and fatty acids present; spores 125-165(-200) µm long; S.Am...... P. anisospora Müll.Arg. 14a. Lichexanthone alone; spores 90-95 µm long; Aus...... P. subcerussata A.W.Archer Spores 125-155 µm long; S.Am ...... P. tetrathalamia var. confirmans Müll.Arg.

15(8a). 2-Chlorolichexanthone present ...... 16 15a. Polychlorinated lichexanthones present ...... 17

16. 2-Chlorolichexanthone with stictic acid present; spores 120-165 µm long; Java...... P. placentiformis (Mont. & Bosch) Müll.Arg. 16a. 2-Chlorolichexanthone with stictic and 2'-O-methylperlatolic acids present; spores 115-125 µm long; Aus ...... P. pallida A.W.Archer & Elix

17. 2,4-, 2,5- Dichlorolichexanthones and 2,4,5-trichlorolichexanthone present ...... 18 17a. 4,5-Dichlorolichexanthone present ...... 20

34

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

18. Thallus isidiate; with stictic acid; spores 80-95 µm long; Aus, NZ ...... P. subisidiosa A.W.Archer 18a. Thallus not isidiate; ...... 19

19. Additional 2-O- and 2'-O-methylperlatolic acids present; ...... spores 80-110 µm long; Aus, PNG, New Caledonia [±stictic acid] .... P. elliptica Müll.Arg. 19a. Additional stictic acid present ; spores 55-75 µm long; Qld...... P. glabra A.W.Archer & Elix

20. 4,5-Dichlorolichexanthone only present; spores 80-110 µm long; Aus...... P. leucophaea Elix & A.W.Archer 20a 4,5-Dichlorolichexanthone and additional compounds present ...... 21

21. 4,5-Dichlorolichexanthone and planaic or confluentic acid present ...... 22 21a. 4,5-Dichlorolichexanthone and stictic acid present ...... 23

22. 4,5-Dichlorolichexanthone and planaic acid present; spores 75-97 µm long; Aus ...... P. leucothelia Müll.Arg. 22a. 4,5-Dichlorolichexanthone and confluentic acid present; spores 74-100 µm; Cook Islands ...... P. rarotongensis A.W.Archer & Elix

23. 4,5-Dichlorolichexanthone and stictic acid present ...... 24 23a. 4,5-Dichlorolichexanthone, stictic acid and additional compounds present ...... 25

24. Spores 75-150 µm long; N.Am, S.Am...... P. tetrathalamia (Fée) Nyl. 24a. Spores 130-150 µm long; West Indies ...... P. ascisidioides Vain

25. 4,5-Dichlorolichexanthone, stictic acid, ±2-chlorolichexanthone ±4-chlorolichexanthone present; spores 100-135 µm long; S.Am...... P. tetrathalamia var. decipiens Müll.Arg. 25a. 4,5-Dichlorolichexanthone, stictic acid and depsides present ...... 26

26. 4,5-Dichlorolichexanthone, stictic and confluentic acids present; spores 105-130 µm; Cook Islands ...... P. rarotongensis var. stictica A.W.Archer & Elix 26a. 4,5-Dichlorolichexanthone, stictic and 2'-O-methylperlatolic acids present; ...... spores 80-106 µm long ...... 27

27. Thallus isidiate; Thailand ...... P. microstoma Müll.Arg. var. isidiata Jariang. 27a. Thallus not isidiate; Aus, Java, PNG, New Caledonia ...... P. microstoma Müll.Arg.

Group 19. Apothecia present, disciform; ascospores 8 per ascus, on rock

1. 4,5-Dichlorolichexanthone and 2-O-methylperlatolic acid present; spores 45-55 µm long; ... on moss; NZ ...... P. scottii Elix & A.W.Archer 1a. Depsidones present ...... 2

2. Fumarprotocetraric or salazinic acid present ...... 3 2a. Norstictic or stictic acid present ...... 4

3. Fumarprotocetraric acid present; spores 23-28 µm long; thallus isidiate; Eur ...... P. oculata (Dickson) Th. Fr. 3a. Salazinic acid present; 24-39 µm long; thallus lacking isidia; Falkland Is...... P. salacinifera Messuti & A.W.Archer

4. Stictic acid present...... 5 4a. Norstictic acid present ...... 6

5. On moss; spores 80-120 µm long; NZ ...... P. tyloplaca Nyl. 5a. On rock; spores 25-42 µm long; Eur, N.Am...... P. chiodectonoides Bagl. ex Massal. 

35 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

6. Spores 18-24 µm long; NZ, S.Am...... P. monticola Messuti & A.W.Archer 6a. Spores 30-40(-47) µm long; S. Hemisphere .. Lepra erubescens (Taylor) A.W.Archer & Elix ...... [P. erubescens (Taylor) Nyl. ]

Group 20. Apothecia present, disciform; ascospores 8 per ascus, on wood or bark

1. Lichen compounds absent ...... 2 1a. Lichen compounds present ...... 4

2. Spores 15-20 µm long; disc black, white pruinose; NZ ...... P. sporellula A.W.Archer & Elix 2a. Spores > 20 µm long ...... 3

3. Spores 34-41 µm long; disc pale grey, epruinose; NZ P. duppensis A.W.Archer & Malcolm 3a. Spores 35 x 19 µm [chemistry unclear] ...... P. diluta C.Björk, G.Thor & T.Wheeler

4. Thiophaninic or picrolichenic acid present...... 5 4a. Depsidones or depsides present ...... 6

5. Thiophaninic acid present; spores 60-110 µm long; Eur, N.Am.P. hymenea (Ach.) Schaer. 5a. Picrolichenic acid present; spores 19-27 µm long; Aus, NZ ...... P. truncata Kremp.

6. Depsidones [fumarprotocetraric, norstictic or psoromic acids] present ...... 7 6a. Depsides present ...... 14

7. Fumarprotcetraric acid present ...... 8 7a. Norstictic or psoromic acid present ...... 11

8. Fumarprotocetraric acid and gyrophoric acid or protocetraric present ...... 9

8a. Fumarprotocetraric acid ± succinicprotocetraric acid present ...... 10

9. Fumarprotocetraric and gyrophoric acid; spores 22-30 µm; N.Am ...... P. suboculata Brodo & Dibben 9a. Fumarprotocetraric and protocetraric acid present; spores 18-20 µm long; ...... Eur, N.Am; also sterile and sorediate ...... P. borealis Erichs.

10. Fumarprotocetraric acid only present;spores 10-14 µm long; Eur . P. pupillaris (Nyl.) Th.Fr. 10a. Fumarprotocetraric and succinicprotocetraric acids present; spores 12-22 µm long; N.Am ...... P. subambigens Dibben

11(7a). Psoromic acid present; spores 27-40 µm long; China .... P. wulingensis Z.T.Zhao & Z.S.Sun 11a. Norstictic acid present ...... 12

12. Norstictic acid and lichexanthone present; spores 16-22 µm long; S.AmP. phlyctaenula Nyl. 12a. Norstictic acid only present; S.Af...... 13

13. Spores 12-22 µm long ...... P. ambigens Nyl. 13a. Spores 20-32 µm long ...... P. elatior Stirt.

14(6a). Lecanoric or gyrophoric acid present ...... 15 14a. Thamnolic acid or 5-O-methylhiascic acid present ...... 18

15. Gyrophoric acid present; spores 20-40 µm long; N. Hemisphere, ...... P. carneopallida (Nyl.) Anzi ex Nyl. 15a. Lecanoric acid present ...... 16

16. Lichexanthone present; spores 1-seriate, 30-47 µm; S.Am ...... P. pycnophora Nyl.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

16a. Lecanoric acid ± gyrophoric acid present ...... 17

17. Spores 58-75 µm long; PNG ...... P. kaindiensis A.W.Archer, Elix & Streimann 17a. Spores 14-22 µm long; with additional gyrophoric acid; N.Am .. P. glaucomela (Tuck.) Nyl.

18. 5-O-Methylhiascic acid ± stictic acid present; spores 33-54 µm long; Alaska ...... P. mccroryae Börk, Goward & Sprib. 18a. Thamnolic acid present ...... 19

19 Spores globular, 20-25 µm diam.; PNG ...... P. gongylospora Elix & A.W.Archer 19a. Spores ellipsoid ...... 20

20. Spores 22-32 µm long; Aus, NZ ...... P. thamnolica A.W.Archer 20a. Spores 30-50 µm long; Malaysia, PNG ...... P. epitheciifera Sipman

Group 21. Apothecia present, verruciform; ascospores 8 per ascus, on rock or moss

1. Xanthones present ...... 2 1a. Xanthones absent ...... 19

2. Lichexanthone or derivatives present ...... 3 2a. Arthothelin, thiophanic or thiophaninic acids present ...... 12

3. Lichexanthone present ...... 4 3a. Chlorolichexanthones present ...... 8

4. Lichexanthone and stictic acid present; spores 2-seriate, 115-150 µm long; Aus, Easter Island ...... P. hadrocarpa Zahlbr.  4a. Lichexanthone and depsides ± protocetraric acid present ...... 5

5. Lichexanthone, planaic and protocetraric acids present; spores rough walled, 1-seriate ...... 80-125 µm long; Central Afr ...... P. lambionii A.W.Archer et al. 5a. Lichexanthone and depsides present ...... 6

6. Lichexanthone and 2-O-methylconfluentic acid present; spores 1-seriate, 60-70 µm long; ... Central Afr...... P. clercii Messuti & A.W.Archer 6a. Lichexanthone and 2-O-methylperlatolic acid present ...... 7

7. On rock; spores 75-105 µm long; Aus, PNG, S.Am, Mexico ...... P. consanguinea Müll.Arg. 7a. On dead moss; spores 80-95 µm long; Japan ...... P. nagasakensis Nyl.

8. 2-Chlorolichexanthone and 2'-O-methylperlatolic acid present; spores 75-90 µm; Brazil...... P. saxatilis A.W.Archer & Elix 8a. 4,5-Dichlorolichexanthone present ...... 9

9. 4,5-Dichlorolichexanthone and 4-chlorolichexanthone present; on moss; spores 2-seriate, 150-185 µm long; S.Am ...... P. confundens Nyl. 9a. 4,5-Dichlorolichexanthone and norstictic acid or 2'-O-methylperlatolic acid present ...... 10

10. Norstictic acid present; spores 70-85 µm long, irreg. 2-seriate; Ins. Rodrigues ...... P. impallescens Nyl. 10a. 2'-O-Methylperlatolic acid present ...... 11

11. Spores 2-seriate, 85-105 µm; Aus ...... P. lavata Müll.Arg. 11a. Spores 1-seriate, 45-75 µm; Aus, NZ ...... P. lophocarpa Körb.

12(2a). Thiophaninic acid present ...... 11 12a. Arthothelin or thiophanic acid present ...... 17

37 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

11. Thiophaninic acid only present; spores 2-seriate, 62-75 µm long; S.Afr...... P. diaziana A.Massal. Spores 65 µm long; Eur, Asia ...... P. flavicans Lamy 11a. Additional depsidones [norstictic or stictic acids] or 2-O-methylperlatolic acid present ..... 12

12. 2-O-Methylperlatolic acid present; spores 1-seriate, 60-85 µm long; ...... Aus, NZ, PNG, Zimbabwe...... P. petrophyes C. Knight 12a. Norstictic or stictic acids present ...... 13

13. Thiophaninic acid and norstictic acid present; spores 85-110 µm long; Eur; ...... P. huneckiana Feige & Lumbsch 13a. Thiophaninic acid and stictic acid present ...... 14

14. Thallus lacking isidia and soredia ...... 15 14a. Thallus isidiate or sorediate ...... 16

15. Spores 2-seriate, 70-90 µm long; Aus ...... P. dissita Elix & A.W.Archer 15a. Spores 1-seriate, 65-80 µm long; S.Am ...... P. hypoxantha Malme 15b. Spores ?seriate, 70-100 µm long; Eur...... P. rupicola (Fr.) Harm.

16. Thallus isidiate; spores 1-seriate, 50-90 µm long; Aus ...... P. xanthodactylina A.W.Archer & Elix 16a. Thallus sorediate; spores 50-75 µm long, 1-seriate; Aus, NZ, PNG ...... P. xanthoplaca Müll.Arg.

17(10a) Thiophanic acid present; spores 2-seriate, 60-85 µm long; Aus ...... P. thula A.W.Archer 17a. Arthothelin present ...... 18

18. Spores subglobose, hyaline to brown, 1-seriate, 12-30 µm long; SW US ...... P. occidentalis Bratt, Lumbsch & Schmitt 18a. Spores ellipsoid, hyaline to pale grey, 2-seriate, 60-75 µm long; Aus, NZ S.Afr. S.Am...... P. melanospora Nyl. 

19(1a) Lichen compounds absent; spores 2-seriate, 37-55 µm long; Aus. .... P. paratropa Müll.Arg. Spores 1-seriate, 50-58 µm long; S.Afr...... P. pseudomelanospora V.Wirth & Elix

19a. Lichen compounds present ...... 20

20. 2'-O-Methylperlatolic acid present ...... 21 20a. Norstictic, stictic or virensic acid present ...... 22

21. Spores hyaline, 1-seriate, 65-85 µm long; S.Am ...... P. cerebrinula Zahlbr. 21a. Spores hyaline to brown, 2-seriate, 55-80 µm long; N.Am ...... P. islandica Bratt, Lumbsch & Schmitt

22. Stictic acid present; spores 2-seriate, 30-36 µm long; Aus ...... P. macra Müll.Arg. Spores 25-45 µm long; Eur...... P. eiteriana Erichs. 22a. Norstictic or virensic acid present ...... 23

23. Virensic acid present; spores 80-125 µm long; S.Am ...... P. rochae Räsänen 23a. Norstictic acid present ...... 24

24. Spores 1-seriate ...... 25 24a. Spores 2-seriate ...... 26

25. Spores 80-135 µm long; ± virensic acid; S.Am ...... P. patagonica Müll.Arg. 25a. Spores 50-70µm long, N.Am, Arctic ...... P. octomela (Norm.) Erichs

26. Spores 115-150 µm long; NZ, S.Am...... P. perrimosa Nyl.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

26a. Spores 87-112 µm long; S.Am ...... P. colorans Malme

Group 22 Apothecia present, verruciform; ascospores 8 per ascus, on wood or bark

1. Xanthones present ...... Subgroup 22A 1a. Xanthones absent ...... Subgroup 22B

Subgroup 22A Xanthones present

1. Lichexanthone or chlorolichexanthones present ...... 2 1a. Norlichexanthone derivatives or thiophaninic acid or its derivatives present ...... 82

2. Lichexanthone present ...... 3 2a. Chlorolichexanthones present ...... 27

3. Lichexanthone only present ...... 4 3a. Lichexanthone with depsides and/or depsidones present ...... 5

4. Spores 2-seriate, 120-140 µm; S.Am ...... P. phaeostoma Müll.Arg. 4a. Spores 1-seriate, 84-100 µm long; Aus ...... P. albula A.W.Archer & Elix

5. Lichexanthone and 2,2'-di-O-methylstenosporic acid ± 2-O-methylstenosporic acid present ...... 6 5a. Lichexanthone and stictic or norstictic or protocetraric acid present ...... 8

6. Lichexanthone and 2,2'-di-O-methylstenosporic acid ± 2-O-methylstenosporic acid present; spores 64-78 µm long; Thailand ...... P. alboaspera var. deficiens Jariang. & A.W.Archer 6a. Lichexanthone and 2,2'-di-O-methylstenosporic acid present; 2-O-methylstenosporic acid absent ...... 7

7. Spores 52 x 27 µm [fide Müll.Arg.]; Central Afr...... P. subareolata Müll.Arg. 7a. Spores 100-125 µm long; Aus ...... P. pinnaculata A.W.Archer & Elix

8. Lichexanthone and norstictic acid ± stictic acid; spores 16-22 µm long; S.Am ...... P. phlyctaenula Nyl. 8a. Lichexanthone with stictic or protocetraric acid ...... 9

9. Lichexanthone and protocetraric acid present ...... 10 9a. Lichexanthone and stictic acid present ...... 13

10. Depsides absent; spores 2-seriate, 70-80 µm long; S.Am ...... P. rhodostomoides Vain. 10a. Depsides [2'-O-methylperlatolic or 2,2'-di-O-methylstenosporic acids] present...... 11

11. 2,2'-Di-O-methylstenosporic acid present; spores 2-seriate, 60-100 µm long; ...... Thailand ...... P. nanensis Jariang. & A.W.Archer 11a. 2'-O-Methylperlatolic acid present ...... 12

12. Spores 1-seriate, 80-125 µm long, rough-walled, with additional planaic acid; ...... Central Afr...... P. lambionii A.W.Archer et al. 12a. Spores 55-60 µm long, smooth-walled; S.Am ...... P. curatellae Malme

13. Lichexanthone and stictic acid present, depsides absent ...... 14 13a. Lichexanthone, stictic acid and depsides present ...... 17

39 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

14. Thallus sorediate or spores rough-walled ...... 15 14a. Thallus esorediate and spores smooth-walled ...... 16

15. Thallus sorediate; spores 98-111 µm long; Central Afr...... P. fosseyae A.W.Archer et al. 15a. Thallus esorediate; spores 125-175 µm long, with rough walls; West Indies ...... P. parnassia Vain.

16. Spores 2-seriate, 100-140 µm long; S.Am, India, Sri Lanka P. dehiscens Müll.Arg. 16a. Spores 1-seriate, 75-90 µm long; S.Am...... P. nigrata Kremp.

17.(13a) Sekikaic, confluentic or ...... 2-O-methylperlatolic acids present ...... 18 17a. Stenosporic acid derivatives present ...... 21

18. Sekikaic or confluentic acids present ...... 19 18a. 2-O-Methylperlatolic acid present ...... 20

19. Sekikaic acid present; spores 2-seriate, 100-140 µm long; Aus ...... P. dehiscens var. sekikaica A.W.Archer & Elix 19a. Confluentic acid present; spores 2-seriate, 87-110 µm long; SAm ...... P. gracilis f. heteromera Müll.Arg.

20. Spores 75-105 µm long; Aus ...... P. leucostigma Müll.Arg. 20a. Spores 60-90 µm long; N.Am ...... P. paratuberculifera Dibben

21. Lichexanthone, stictic and 2'-O-methylstenosporic acids present ...... 22 21a. Lichexanthone, stictic and 2,2'-di-O-methylstenosporic acids present ...... 23

22. Spores 2-seriate, 70-110 µm long; Thailand ...... P. platycarpa Jariang. 22a. Spores 1-seriate, 56-62 µm long; Aus...... P. montoensis A.W.Archer & Elix

23. Spores 1-seriate ...... 24 23a. Spores 2-seriate ...... 25

24. Spores 50-67 µm long; Brazil ...... P. verruculifera Vain. 24a. Spores 60-84 µm long; Colombia ...... P. leioplaca var. pycnocarpa Nyl.

25. Ostioles black; spores 62-75 µm long; S.Afr ...... P. congesta Vain. 25a. Ostioles pale ...... 26

26. Spores 90-100 µm long; Aus...... P. alboaspera A.W.Archer & Elix 26a. Spores 75-85 µm long; S.Afr...... P. glaucocinerea Vain.

27(2a) 4,5-Dichlorolichexanthone present ...... 28 27a. 2-Chlorolichexanthone or 2,4- and 2,5-dichlorolichexanthones and ...... 2,4,5-trichlorolichexanthone present ...... 63

28. 4.5-Dichlorolichexanthone only present ...... 29 28a. 4.5-Dichlorolichexanthone and depsidones and/or depsides present ...... 34

29. Spores 150-200 µm long; [± 4-chlorolichexanthone]; S.Am ...... P. confundens Nyl. 29a. Spores <100 µm long ...... 30

30. Spores 1-seriate, 30-37 µm long; NZ ...... P. parvula A.W.Archer & Elix 30a. Spores 2-seriate, >50 µm long ...... 31

31-. Spores 60-90 µm long ...... 32

40

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

31a. Spores 54-70 mm long ...... 33

32. Spores 8/ascus; Aus, Asia, Central Afr...... P. rigida Müll.Arg. 32a. Spores 6/ascus [also ± stictic acid]; Finland...... P. raesaenenii Erichs.

33. Spores 55-70 µm long; S.Am; ostioles conspicuous, black, 1-4, in translucent zones ...... P. mundula Müll.Arg. 33a. Spores 54-60 µm long; China; ostioles 2-4, black, punctiform ...... P. subrosaceae var. octospora Zahlbr.

34(28a). Depsidones [norstictic or stictic acids] present; depsides and 9-O-methylalternariol absent ...... 35 34a. Depsides or 9-O-methylalternariol present, ±stictic acid ...... 45

35. 4,5-Dichlorolichexanthone and norstictic acid present; spores 2-seriate, 67-85 µm long; Aus ...... P. undulata Müll.Arg. 35a. 4,5-Dichlorolichexanthone and stictic acid present ...... 36

36. Spores 1-seriate ...... 37 36a. Spores 2-seriate ...... 42

37. Species circumboreal ...... 38 37a. Species in Southern Hemisphere ...... 41

38. Species predominantly circumboreal ...... 39 38a. Species predominantly European ...... 40

39. Spores 18-46 µm long; ...... P. sommerfeltii (Flörke) Fr. 39a. Spores 40-70 µm long; also in Japan ...... P. alpina Hepp ex Ahles

40. Spores 30-60 µm long; S Eur...... P. constricta Erichs. 40a. Spores 60-90 µm long; central Eur ...... P. caesioumbrina Eitner

41. Spores 65-70 µm long; ostioles white; Aus ...... P. leucostomoides Zahlbr. 41a. Spores 48-72 µm long; ostioles black; S.Am ...... P. melanostoma Kremp.

42(36a). Spores 55-80 µm long; Aus, PNG, China, Japan ...... P. leiocarpella Müll.Arg. 42a. Spores > 100 µm long ...... 43

43. Spores 180-270 µm long; Japan ...... P. akagiensis Vain. 43a. Spores < 150 µm long ...... 44

44. Spores 100-130 µm long; West Indies ...... P. polysticta Vain. 44a. Spores 130-140 µm long; Japan ...... P. laeviganda Nyl.

45. 4.5-Dichlorolichexanthone, stictic acid and stenosporic acid derivatives present..... 46 45a. 4.5-Dichlorolichexanthone and depsides or 9-O-methylalternariol present; stictic acid absent ...... 47

46. 2-O-Methylstenosporic acid present; spores 2-seriate, 80-94 µm long; Thailand ...... P. kansriae var. stictica Jariang. 46a. 2,2'-Di-O-methylstenosporic and 2,2'-di-O-methyldivaricatic acids present; ...... spores 64-102 µm long; Thailand...... P. subplanaica var. stictica Jariang.

47. 9-O-Methylalternariol present; spores 2-seriate, 75-87 µm long; PNG ...... P. praecipua A.W.Archer & Elix 47a. Perlatolic or stenosporic acid derivatives present ...... 48

48. Stenosporic acid derivatives present ...... 49 48a. Perlatolic acid derivatives or confluentic, miriquidic or divaricatic acids present

41 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

...... 52

49. O-Methylstenosporic acid derivatives present ...... 50 49a. 2,2'-Di-O-methylstenosporic acid present ...... 51

50. 2-O-Methylstenosporic acid present; spores 2 seriate, 64-80 µm long; India ...... P. rimosa Awasti & Srivastava 50a. 2'-O-Methylstenosporic acid present; spores 1 seriate, 70-80 µm long; Aus ...... P. praetermissa A.W.Archer & Elix

51. Spores 1-seriate, 70-110 µm long; Aus, NZ; PNG ...... P. subplanaica A.W.Archer & Elix 51a. Spores 2-seriate, 60-78 µm long; Thailand ...... P. boklueaensis Jariang.

52. Miriquidic or confluentic acid present ...... 53 52a. Perlatolic acid, perlatolic acid derivatives or divaricatic acid present ...... 55

53. Miriquidic acid present; spores 1-seriate, 75-95 µm long; Aus ...... P. novaehollandiae A.W.Archer 53a. Confluentic acid present ...... 54

54. Confluentic acid only present; spores 1-seriate, 60-75 µm long; PNG ...... P. weberi A.W.Archer & Elix 54b. Confluentic and 2'-O-methylperlatolic acids present; spores 50-60 µm long; Eur...... P. werneriana Boqueras

55. Perlatolic acid or divaricatic acid present ...... 56 55a. 2'-O-Methylperlatolic, 2-O-methylperlatolic or planaic acid present ...... 57

56. Perlatolic acid present; spores 1-seriate, 50-60 µm long; Aus ......

...... P. minispora A.W.Archer & Elix 56a. Divaricatic acid present; spores 2-seriate, 75-82 µm long; Aus ...... P. pseudothwaitesii A.W.Archer & Elix

57. 2'-O-Methylperlatolic or 2-O-methylperlatolic acid present ...... 58 57a. Planaic acid present ...... 61

58. 2-O-Methylperlatolic acid and methyl 2-O-methylperlatolate present; ...... spores 1-seriate, 50-60 µm long; Aus...... P. xenismota A.W.Archer & Elix 58a. 2'-O-Methylperlatolic acid present ...... 59

59. Spores 2-seriate, 55-110 µm long; Aus, PNG, Sri Lanka, N.Am ...... P. mesotropa Müll.Arg. 59a. Spores 1-seriate ...... 60

60. Spores 32-65 µm long; Aus...... P. pertractata Stirt. 60a. Spores 55-65 µm long; with additional 2,4,5-trichlorolichexanthone; PNG ...... P. allogiberosa A.W.Archer & Elix

61. Planaic acid and 2-O-methylperlatolic acid present; spores 45-55 µm long; Aus...... P. placocarpa A.W.Archer & Elix 61a. Planaic acid only present ...... 62

62. Spores 80-100 µm; E.Aus ...... P. planaica A.W.Archer & Elix 62a. Spores 40-65 µm; W.Aus ...... P. subarida A.W.Archer & Elix

63(27a) 2-Chlorolichexanthone present; dichloro- and trichlorolichexanthones absent ...... 64 63a. 2,4- and 2,5-Dichlorolichexanthone and 2,4,5-trichlorolichexanthone present ...... 68

42

Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

64. 2-Chlorolichexanthone only present; spores 45-50 µm long; S.Am ...... P. torquatella Müll.Arg. 64a. 2-Chlorolichexanthone and depsides or depsidones present ...... 65

65. 2-Chlorolichexanthone and depsidones [stictic, constictic acids] present ...... 66 65a. 2-Chlorolichexanthone and depsides [divaricatic, 2'-O-methylstenosporic acids] present 67

66. Stictic acid (major) and constictic acid (minor) present; spores...... 65-80 µm long; S.Afr ...... P. cryptostoma Müll.Arg. 66a. Constictic acid (major) and stictic acid (minor) present; spores ...... 55-66 µm long; S.Am ...... P. delicatula Müll.Arg.

67. 2'-O-Methylstenosporic acid present; spores 1-seriate, 60-70 µm long: Aus ...... P. boweniana A.W.Archer & Elix 67a. Divaricatic acid present; spores 2-seriate, 70-92 µm long: Aus ...... P. orarensis A.W.Archer & Elix

68(63a). Chloroxanthones only present ...... 69 68a. Chloroxanthones and stictic acid and/or depsides present ...... 73

69. Spores 105-135 µm long; rough-walled; PNG ...... P. anomalospora A.W.Archer, Elix & Streimann 69a. Spores <100 µm long ...... 70

70. Spores 50-75 µm long ...... 71 70a. Spores 40-100 µm long ...... 72

71. Spores 62-75 µm long; ostioles brown, conspicuous; S.Am ...... P. paraguayensis Müll.Arg. 71a. Spores 48-60 µm long; ostioles pale grey, translucent; West Indies ...... P. glaucopunctata Vain.

72a. Spores 70-100 µm long; N.Am ...... P. ostiolata Dibben 72b. Spores 40-54 µm long: S.Am...... P. ferax (Müll.Arg.) A.W.Archer & Elix ...... [basionym: P. torquatella var. ferax Müll.Arg.]

73(68a). Stictic acid present ...... 74 73a. Stictic acid absent ...... 75

74. Spores 2-seriate, 50-75 µm long; Aus ...... P. lordhowensis A.W.Archer & Elix; 74a. Spores 1-seriate, 45-50 µm long; additional 2-O-methylperlatolic acid present; Aus ...... P. gundermanica A.W.Archer & Elix,

75. Spores 1-seriate, 45-50 µm long; 2-O-Methyl- and 2'-O-methylperlatolic acids present; Aus ...... P. complanata A.W.Archer & Elix 75a. Spores 2-seriate, 85-112 µm long; 2-O-methylperlatolic acid present; S.Am...... [syntype, leg. Schenk 1891] ...... P. "polita" Müll.Arg

76(1a). Arthothelin, 6-O-methylarthothelin or thiophanic acid present ...... 77 76a. Thiophaninic acid or 4-chloro-6-O-methylnorlichexanthone present ...... 82

77. Arthothelin and thiophanic acid present; spores ?-seriate ...... 78 77a. 6-O-Methylarthothelin and/or arthothelin present; spores 2-seriate ...... 79

78. Spores 60-75 µm long; NZ ...... P. bartlettii A.W.Archer & Elix 78a. Spores 80-95µm long; PNG ...... P. karkarensis A.W.Archer & Elix

79. Spores 100-125 µm long; S.Am ...... P. papillulata Nyl.

43 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

79a. Spores <100 µm long ...... 80

80. Spores 60-87 µm long; 6-O-methylarthothelin and arthothelin; S.Am, Aus, PNG ...... P. oblongata Müll.Arg. 80a. Spores 55-80 µm long; with 6-O-methylarthothelin or arthothelin ...... 81

81. Spores 55-60 µm; 6-O-methylarthothelin only; S.Am...... P. candida Müll.Arg. 81a. Spores 56-80 µm long; arthothelin only; N.Am...... P. appalachensis Lendemer

82(76a). 4-Chloro-6-O-methylnorlichexanthone present; spores 1-seriate, 70-87 µm long; Sri Lanka ...... P. subdepressa Müll.Arg. 82a. Thiophaninic acid present ...... 83

83. Thiophaninic acid only present ...... 84 83a. Thiophaninic acid and depsides and/or depsidones present ...... 90

84. Spores 1-seriate, 50-70 µm long; ostioles black; NZ ...... P. laevis C.Knight 84a. Spores 2-seriate ...... 85

85. Spores 85-115 µm long; ostioles black; NZ ...... P. theochroa Kremp. 85a. Spores 50-95 µm long ...... 86

86. Spores 75-95 µm long; ostioles brown; Aus ...... P. thiophaninica A.W.Archer 86a. Spores ≤ 70 µm long ...... 87

87. French Antilles and Eur...... 88 87a. New Caledonia and S.Am...... 89

88. Spores 52-65 µm long; ?-seriate; French Antilles ...... P. flavens Nyl. 88a. Spores 30-70 µm long; 1-2 seriate; Eur...... P. plombii B.deLesd.

89. Spores 8/ascus; S.Am ...... P. meridionalis var ochrostoma Müll.Arg. 89a. Spores 4 or 8? /ascus; New Caledonia ...... P. endochroma Müll.Arg.

90(83a).Depsidones ± depsides present ...... 91 90a. Depsides present, stictic acid absent ...... 104

91. Depsidones [norstictic, protocetraric, stictic, virensic or variolaric acids] present; ...... depsides absent ...... 92 91a. Stictic acid and depsides present ...... 109

92. Norstictic or protocetraric acid present ...... 93 92a. Stictic or variolaric or virensic acid present ...... 95

93. Norstictic acid present: Spores 55-110 µm long; N.Am ...... P. rubefacta Erichs. [Spores 52-68 µm; N.Africa ...... P. praelutescens Werner (nom. inval.)] 93a. Protocetraric acid present ...... 94

94. Protocetraric acid present, stictic acid absent; spores 1-seriate, 55-85 µm long; S.Am ...... P. glaziovii Müll.Arg. 94a. Protocetraric and stictic acids present; spores 1-seriate, 50-62 µm long; S.Am ...... P. lignicola Malme

95(92a).Stictic acid present ...... 96 95a. Variolaric or virensic acid present ...... 103

96. Thallus sorediate or isidiate...... 97 96a. Thallus lacking soredia and isidia ...... 98

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

97. Thallus sorediate; spores 2-seriate, 60-65 µm long; Aus ...... P. xanthosorediata A.W.Archer 97a. Thallus isidiate; spores 1-seriate, 60-100 µm long; widespread; ± 2'-O-methylperlatolic acid; rarely fertile ...... P. flavida (DC.) Laundon

98. Additional constictic acid present; spores 1-seriate ...... 99 98a. Additional constictic acid absent; spores 1- or 2-seriate ...... 100

99. Spores 50-75 µm long; S.Am ...... P. rhodostoma Nyl. 99a. Spores 70 x 28 µm [fide Müll.Arg.]; E.Afr...... P. cinctula Müll.Arg.

100. Spores 1-seriate ...... 101 100a. Spores 2-seriate ...... 102

101. Spores 52-72 µm long; S.Hemisphere...... P. leioplacella Nyl. 101a. Spores 48-80 µm long; Japan ...... P. obsolescens Nyl.

102. Spores 50-80 µm long; ostioles not mammiform; N.Am, Aus, Seychelles ...... P. texana Müll.Arg. 102a. Spores 56-80 µm long; ostioles mammiform, pale yellow; E.Afr...... P. xanthothelia Müll.Arg.

103(95a). Variolaric acid present; spores 65-90 µm long; N.Am ..... P. epixantha R.C.Harris 103a. Virensic acid present; spores 60-70 µm long; Aus., N.Am ..... P. virensica R.C.Harris

104(90a). Gyrophoric acid present; spores 2-seriate, 60-130 µm long; N.Am, Eur ...... P. hymenea (Ach.) Schaer. 104a. Perlatolic or stenosporic acid derivatives present ...... 105

105. 2-O-Methylstenosporic acid present; spores 2-seriate, 70-80 µm long; S.Am...... P. araucariae Müll.Arg. 105a. Perlatolic acid or its derivatives present ...... 106

106. Perlatolic acid present; spores 1-seriate, 60-70 µm long ...... 107 106a. 2'-O-Methylperlatolic or 2-O-methylperlatolic acid present ...... 108

107. Apothecia scattered; Aus ...... P. injuneana A.W.Archer & Elix 107a. Apothecia confluent; Korea ...... P. jogyeensis J.S.Park & J.-S.Hur

108. 2'-O-Methylperlatolic acid present; spores 2-seriate, 40-80 µm long; Argentina ...... P. formosensis Messuti & A.W.Archer 108a. 2-O-Methylperlatolic acid present; spores 1-seriate, 60-82 µm long; Aus ...... P. xylophyes A.W.Archer [Spores 52-72 µm long, 1-seriate; India; 2-O-methylperlatolic acid and traces ...... thiophaninic acid present; India ...... ?syn: P. amarkantakana Srivastava & Awasthi]

109(91a)Spores 2-seriate, 64-90 µm long;; perlatolic and hyperlatolic acids present; ...... Thailand ...... P. hylocola Jariang. & A.W.Archer 109a Spores 1-seriate ...... 110

110. 4-O-Methylisocryptochlorophaeic acid present; spores 75-100 µm long; Aus ...... P. paradoxica A.W.Archer & Elix 110a. 2,2'-Di-O-methylstenosporic acid present; spores 80-105 µm long; Aus...... P. dharugensis A.W.Archer & Elix

Subgroup 22B Xanthones absent

1a. Lichen compounds absent ...... 2 1a. Lichen compounds present ...... 13

45 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

2. Spores 2-seriate ...... 3 2a. Spores 1-seriate ...... 8

3. Spores ≥70 µm long ...... 4 3a. Spores ≤ 70 µm long ...... 5

4. Spores 120-150 µm; rough; Cook Islands ...... P. megacarpa A.W.Archer & Elix 4a. Spores 70-110 µm long; smooth; Aus., S.Am, Cent. Afr...... P. subrigida Müll.Arg.

5. Spores 35-55 µm; China ...... P. subochracea Stirton 5a. Spores 45-70 µm long ...... 6

6. Spores 48-70 µm long; ostioles red-black; NZ, Chile ...... P. leucodes C. Knight 6a. Spores 45-56 µm long ...... 7

7. Spores 45-50 µm long; ostioles small, black, conspicuous; S.Am ...... P. melaleuca var. octospora Müll.Arg.

7a. Spores 45-56 µm long; ostioles large, black, conspicuous; ?N.Am...... P. gonolobina Müll.Arg.

8. Spores 70-105 µm long ...... 9 8a. Spores 50-75 µm long ...... 10

9. Spores 70-100 µm long; ostioles conspicuous, black; NZ ...... P. micropora Kremp. 9a. Spores 75-105 µm long; ostioles inconspicuous, black; S.Am ... P. dilatata Müll.Arg.

10. Apothecia flattened ...... 11 10a. Apothecia flattened hemispherical ...... 12

11. Ostioles black, inconspicuous; S.Am, S.Afr, NZ ...... P. cryptocarpoides Vain. 11a. Ostioles conspicuous, black; S.Am ...... P. brasiliana Zahlbr.

12. Ostioles translucent; NZ ...... P. albissima Müll.Arg. 12a. Ostioles conspicuous, black; S.Am ...... P. acromelana Müll.Arg.

13(1a). Depsidones and depsides present ...... 14 13a. Depsides present ...... 35

14. Depsidones [stictic, constictic, norstictic, salazinic, protocetraric] present, depsides absent ...... 15 14a. Stictic acid and depsides present ...... 31

15. Stictic, constictic or norstictic acid present ...... 16 15a. Salazinic or protocetraric acid present ...... 17

16. Stictic or constictic acid present; spores 2-seriate ...... 18 16a. Norstictic acid present ...... 22

17. Salazinic acid present; spores 2-seriate, 90-120 µm long; NZ ...... P. celata A.W.Archer & Elix 17a. Protocetraric acid present; spores 2-seriate, 70-95 µm long; Aus ...... P. errinundrensis A.W.Archer

18. Constictic acid present; spores 80 x 30 µm [fide Müll.Arg.]; E. Afr...... P. aspera Müll.Arg. 18a. Stictic acid present ...... 19

19. Stictic and 5-O-methylhiasic acids present; spores 43 x 18 µm; US ......

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

...... P. mccroryae C.R. Björk, Goward & T. Sprib. 19a. Stictic acid alone present ...... 20

20. Spores 1-seriate, 67-84 µm long; WAfr ...... P. guinea-bissauensis, sp. nov. 20a. Spores 2-seriate ...... 21

21. Spores 37-48 µm long; Marquesas Is...... P. dermatodes Nyl. 21a. Spores 145-170 µm long; Guadalupe ...... P. gibberosula Nyl.

22(16a)Spores 2-seriate ...... 23 22a. Spores 1-seriate ...... 26

23. Spores 180-210 µm long; PNG ...... P. sipmanii A.W.Archer & Elix 23a. Spores <150 µm ...... 23

24. Spores 45-55 µm long; Norfolk Is ...... P. norfolkensis A.W.Archer 24a. Spores >55 µ long ...... 25

25. Spores 80-105 µm; ostioles becoming pseudolecanorine; N.Am, S.Am...... P. propinqua Müll.Arg. 25a. Spores 75-88 µm long; India, China ...... P. himalayensis Awasti & Srivastava

26. Spores 95-115 µm long; S.Am...... P. patagonica Müll.Arg. 26a. Spores <100 µm long ...... 27

27. Spores 17-20 µm diam; [?not Pertusaria]; Norfolk Is ..... P. globospora A.W.Archer, 27a. Spores 50-95 µm long; ...... 28

28. Spores 60-95 µm long ...... 29 28a. Spores 50-75 mm long ...... 30

29. Spores 60-95 µm long; ostioles inconspicuous; E.Afr ...... P. endoxantha Vain. 29a Spores 60 µm long; ostioles, black, conspicuous; W.Afr...... P. personata Müll.Arg.

30. Spores 50-70 µm long; Arctic ...... P. octomela (Norm.) Erichs. 30a. Spores 50-75 µm long; India; +unidentified spot ?depsideP. maculata Singh & Sinha

31(14a). 2,2'-Di-O-methylstenosporic acid present; spores 2-seriate, 68-85 µm long; Thailand ...... P. alboaspera var. disflavens Jariang. 31a. Perlatolic acid derivatives or divaricatic acid present ...... 32

32. Divaricatic acid present; spores 2-seriate, 75-100 µm long; Aus ...... P. sydneyensis A.W.Archer & Elix 32a. Perlatolic acid derivatives present ...... 33

33. 2-O-Methylsuperlatolic acid present; spores 1-seriate, 60-100 µm long; Thailand ...... P. krabiensis Jariang. 33a. 2'-O-Methylperlatolic acid present ...... 34

34. Spores 1-seriate, 74-110 µm long; Thailand ...... P. phulhuangensis Jariang. 34a. Spores 2-seriate, 70-80 µm long; Java ...... P. minor Müll.Arg.

35(13a). Stenosporic acid derivatives present ...... 36 35a. Perlatolic acid or other compounds present ...... 38

36. 2,2'-Di-O-methylstenosporic acid present; spores 2-seriate, 100-120 µm long; S.Am . [cf. syntype with chloroxanthones, supra] ...... P. polita Müll.Arg. 36a. 2-O-Methylstenosporic acid present ...... 37

47 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

37. Spores 1-seriate, 60-75 µm long; PNG ...... P. kagamugana Elix & A.W.Archer 37a. Spores 2-seriate, 70-90 µm long; Aus, S.Am, NZ, India ... P. leucoplaca Müll.Arg.

38. Gyrophoric or divaricatic acid present; spores 2-seriate ...... 39 38a. Perlatolic acid derivatives present ...... 41

39. Gyrophoric acid present; spores 40-64 µm; N.Am ...... P. papillata (Ach.) Tuck 39a. Divaricatic acid present ...... 40

40. Spores 65-75µm; ostioles inconspicuous, pale brown, translucent; PNG ...... P. papuana A.W.Archer & Elix 40a. Spores 56-85 µm; ostioles conspicuous, black; Sri Lanka ...... P. melastomella Nyl.

41. Confluentic acid present ...... 42 41a. Perlatolic acid derivatives present; spores 2-seriate ...... 43

42. Confluentic acid and atranorin present; spores 2-seriate, 50-60 µm long ...... PNG...... P. gorokorana Elix & A.W.Archer 42a. Confluentic and 2'-O-methylperlatolic acids present; spores 2-seriate, 40-50 µm long: S.Am ...... P. nana Müll.Arg.

43. Glomellerific acid present; spores 80-90 µm long; S.Am ...... P. corrugata Kremp. 43a. 2'-O-Methylperlatolic or 2-O-methylperlatolic acid present ...... 44

44. 2'-O-Methylperlatolic acid present ...... 45 44a. 2-O-Methylperlatolic acid present ...... 47

45. 2'-O-Methylperlatolic acid present with additional stictic acid; spores 92-120 µm long; S.Am ...... P. ampliata Erichs. 45a. 2'-O-Methylperlatolic acid present alone ...... 46

46. Spores 62-90 µm long; S.Am...... P. cerussata Malme 46a. Spores 85-102 µm long; Central Afr...... P. kigomensis Dodge

47. 2-O-Methylperlatolic acid present; spores 92-120 µm long; S.Am ...... P. tuberculifera Nyl. 47a. Spores ≤ 100 µm long ...... 47

48. Spores 75-100 µm long; S.Am ...... P. achroiza Nyl. 48a. Spores 65-85 µm long; S.Am; Aus ...... P. mattogrossensis Malme

SYNONYMS

P. akagiensis Vain. syn: P. conformis Vain.

P. alaskensis Erichs syn: P. aleutensis Erichs.

P. albescens (Huds.) Choisy & Werner syn: P. globulifera (Turner) Massal. syn: P. discoidea (Pers.) Malme syn: P. orbiculata (Schreber) Zahlbr. syn: P. tuberculata (Erichs.) Erichs syn: P. deschatresii Werner syn: P. henrici Erichs. ?syn: var. corallina (Zahlbr.) Laundon

P. allanii Zahlbr. syn: P. aucklandica Zahlbr. nom. nud.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. alpina Hepp ex Ahles syn: P. laevigata (Th.Fr.) Anzi [non P. laevigata Müll.Arg.]

P. amara (Ach.) Nyl. syn: P. slesvicensis Erichs syn: P. slesvicensis var. levior Erichs. syn: P. pulvinata Erichs. syn: P. amara var. verrucigera Erichs. syn: P. amara var. macrosora Erichs. syn: P. slesvicensis var. deformis Erichs. syn: P. commutata Müll.Arg. sensu Oshio

P. amarescens Nyl syn: P. flavicans Lamy syn: P. flavosulphurea Vain. syn: P. affinis Erichs. syn: P. leucosoroides Nyl.

P. amaroides H. Magn. syn: P. amara var. tucumanensis Räs.

P. ampliata Erichs. syn: P. tuberculifera var. virens Müll.Arg. syn: P. tetrathalamia var. virens(Müll.Arg.) Zahlbr. P. anisospora Müll.Arg. syn: var. subflavens Müll.Arg. syn: var. major Müll.Arg.

P. antillarum Vain. syn: P. intermedia Vain.

P. areolata (Ach.) A. Massal. syn: P. chionea DC. in Lam. & DC.

P. brasiliana Zahlbr. syn: P. laevigata Müll.Arg. nom. inval., non P. laevigata (Th.Fr.) Anzi

P. carneola (Eschw.) Müll.Arg. syn: P. communis var. carneola Eschw.

P. carneopallida (Nyl.) Anzi ex Nyl. syn: P. protuberans (Sommerf.) Th. Fr. ?syn: P. perfida Nyl. sensu Oshio syn: P. leptocarpa Anzi syn: P. phlyctidia Norm.

P. cerebrinula Zahlbr. syn: P. alta Zahlbr syn: P. elatior Müll.Arg. nom.inval, non P. elatior Stirt.

P. cerussata Malme syn: P. cerussata var. dispersa Malme

P. ceylonica Müll.Arg. syn:P. blumeana Müll.Arg. syn: P. impressula Müll.Arg. syn: P. trisperma Müll.Arg.

49 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. chiodectonoides Bagl. ex Massal. syn: P. nolens Nyl. syn: P. inquinata (Ach.) Th.Fr.

P. cicatricosa Müll.Arg. ?syn: P. communis var. neocaledonica Nyl. syn: P. subtruncata Müll.Arg. ?syn: P. goniostoma Müll.Arg. syn: P. schiffneri Zahlbr.

P. circumcincta Stirt. syn: P. creberrima Stirt. syn: P. krempelhuberi Müll.Arg. syn: P. subverrucosa Kremp. nom. inval., non. Nylander

P. clarkeana A.W.Archer syn: P. confusa A.W.Archer, nom. inval. non. Zahlbr

P. coccodes (Ach.) Nyl. syn: P. polycarpiza Choisy & Werner syn: P. coccodes (Ach.) Nyl. var. petraea Erichs.

P. commutata Müll.Arg. syn: P. vaginatula Vain. syn: P. copiosa Erichs.

P. composita Zahlbr. syn; P. petrophyes Knight var. hokiensis Vain.

P. consanguinea Müll.Arg. syn: P. rudis Müll.Arg. syn: P. mariae de Lesd.

P. corallina (L.) Arn. syn: P. chionea DC syn: P. dubia Nyl. syn: P. communis f. sorediata Nyl. ?syn: P. microthelia Vain. [K++ yellow; SW Africa]

P. coriacea (Th.Fr.) Th.Fr. syn: P. obducens Nyl.

P. coronata (Ach.) Th.Fr. syn: P. isidiifera Erichs. syn: P. subdubia Nyl. var. hungarica Erichs.

P. cribellata Branth syn: P. solitaria H. Magn. nom. inval. non P. solitaria Darb.

P. cryptocarpoides Vain syn: P. spilota A.W.Archer & Malcolm

P. cryptostoma Müll.Arg. syn: P. limbata Vain. syn P. spaniostoma Vain. syn: P. simplex Vain. syn: P. robsonii Dodge

P. dealbata (Ach.) Cromb.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

syn: P. dealbescens Erichs.

P. dehiscens Müll.Arg. syn: P. dehiscens Müll.Arg. syn: P. dehiscens var. alba Müll.Arg. syn: P. dehiscens var. depressior Müll.Arg. syn: P. sulphurescens Müll.Arg. syn: P. major Müll.Arg. syn: P. leioplaca (Ach.) Shaer. var. turgida Müll.Arg. syn: P. splendens Awasti & Srivastava

P. depressa (Fée) Müll.Arg. syn: P. modesta Müll.Arg. syn: P. leioplaca (Ach.) DC in Lamy & DC var. depressula Müll.Arg. syn: P. melaleuca (Smith & Sow.) Duby var. tetramera Müll.Arg. ?syn: P. sinusmexicani Dibben [chemistry unclear]

P. dispersa Vain. syn: P. multiplicans Vain.

P. eiteriana Erichs. syn: P. polycarpa Eitner; nom. inval. non Krempel.

P. endoxantha Vain. syn: P. norstictica A.W.Archer syn: P. macrostomoides Dodge syn: P. prolifera Dodge syn: P. sanguinescens Zahlbr.

P. erubescens (Taylor) Nyl. syn: P. perfida Nyl. syn: P. vepallida Nyl. syn: P. concava Müll.Arg. syn: P. solitaria Darb. syn: P. darbishireana Zahlbr. syn: P. corrugata Darb. [nom. inval. non P. corrugata Kremp.]

P. excludens Nyl. syn: P. inopinata Erichs.

P. flavida (DC.) J.R. Laundon syn: P. lutesecens (Hoffm.) Lamy syn: P. lutescens f. fagicola Høeg

P. glaziovii Müll.Arg. syn: P. subsulphurescens Malme

P. glomerata (Ach.) Schaer. syn: P. diffusilis Erichs.

P. graphica C. Knight syn: P. graphica f. tardiva Zahlbr. syn: P. dinota Stirt. syn: P. thomsoniana Zahlbr.

P. hadrocarpa Zahlbr. syn: P. asperata A.W.Archer

51 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. hartmannii Müll.Arg. syn: P. trypetheliiformis (Nyl.) Nyl. var. hartmannii (Müll.Arg.) Müll.Arg. syn: P. muelleriana Zahlbr. syn: P. albinea Müll.Arg., nom. inval. non P. albinea Tuck.

P. hemisphaerica (Flörke) Erichs. syn: P. speciosa Høeg

P. heterochroa (Müll.Arg.) Erichs. syn: P. pustulata (Ach.) Duby var. heterochroa Müll.Arg. syn: P. maximiliana Klem.

P. hossei (Räs.) A.W.Archer & Osorio syn: P. reducta Stirt. var hossei Räs

P. huangshanensis S.Yu & J.Wu ex Q. Ren cf. P. variolosa (Kremp.) Vain. sensu Singh &Sinha [but spores 80-90 µm] cf. P. sphaeophora Oshio sensu Zhao et al. [spores 167-204 µm]

P. hymenea (Ach.) Schaerer syn: P. wulfenii DC. syn: P. lecanorodes Erichs.

P. impallescens Nyl. ?syn: P. chapadensis Malme

P. irregularis Müll.Arg. non P. irregularis Magn.

P. knightiana Müll.Arg. syn: P. ceuthocarpa [var.] crenulata Stirt. syn: P. whinrayii A.W.Archer

P. laeviganda Nyl syn: P. deplanata Müll.Arg. syn: P. deplanata var. ferruginea Müll.Arg. syn: P. kotsukensis Vain. syn: P. laeviganda Nyl. var. obscurior Vain. syn: P. cinereo-obscurata Vain. syn: P. inabensis Vain.

P. leiocarpella Müll.Arg. syn: P. stenostoma Vain.

P. leioplacella Nyl. syn: P. amblygona Müll.Arg. syn:P. confluens Müll.Arg. syn: P. mesoxantha Müll.Arg.

P. leucodes C. Knight syn: P. cretacea Müll.Arg. syn: P. fernandeziana Messuti

P. leucoplaca Müll.Arg. syn: P. acuta Müll.Arg.

P. leucosora Nyl. syn?; P. albescens var. subflotowiana Erichs. syn?: P. soralifera (J. Steiner) Erichs.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. leucostoma (Bernh.) Massal. syn: P. leioplaca (Ach.) DC syn: P. leucostoma var. dehiscens Erichs. syn: P. plena Anzi syn: P. tabuliformis Erichs. ?syn: sub "P. quassiae" H-NYL 3699 pm pp ex Sri Lanka

P. leucostomoides Zahlbr. syn: P. leucostoma Müll.Arg. [nom inval. non P. leucostoma (Bernh.) Mass.]

P. lophocarpa Körber syn: P. glebosa Müll.Arg. syn: P. arenacea Müll.Arg. syn: P. diffracta Müll.Arg. syn: P. superba Zahlbr. syn: P. dunedina Zahlbr.

P. malmeii Elix & A.W.Archer syn: P. quassiae (Fée) Nyl. var. sordida Malme

P. melaleucoides Müll.Arg. syn: P.galactina Zahlbr. syn: P. atropunctata A.W.Archer

P. melanospora Nyl. syn: P. crassilabra Müll.Arg. syn: P. amaurospora Hellbom

P. meridionalis var ochrostoma Müll.Arg. syn: P. meridionalis var cinerascens Müll.Arg. syn: P. ochrostoma (Müll.Arg.) Vain. [nom. inval. non P. ochrostoma Müll.Arg.]

P. mesotropa Müll.Arg. syn: P. paragibberosa A.W.Archer

P. miscella A.W.Archer syn: P. velatoides A.L. Sm. p.p. P. melanospora Nyl.

P. multipuncta (Turner) Nyl. syn: P. multipuncta var. macrospora A.L. Sm. syn: P. sorediata (Fr.) Fr. syn: P. faginea (L.) Tuck.

P. neilgherrensis (Müll.Arg.) Awasthi & Srivastava syn: P. leioplacoides var. neilgherrensis Müll.Arg.

P. nakamurae (Räsänen) Dibben syn: P. tuberculifera Nyl. var. nakamurae Räs.

P. neolecanina Lumbsch & Nash syn: P. alloluteola A.W.Archer & Elix

P. nigrata Kremp. syn: P. tuberculifera var. ferax Müll.Arg.

53 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. nigrodisca Oshio syn: P. sphaerophora Oshio

P. novaezelandiae Szatala ?syn: P. bengalensis Vain.

P. oblongata Müll.Arg. syn: P. howeana A.W.Archer & Elix

P. ochrostoma Müll.Arg. syn: P. ochrostomoides Vainio nom. superfl.

P. oshioi Wei syn: P. oshioi Dibben, nom. inval. syn: P. macrospora Oshio, nom. inval.

P. otagoana Galloway syn: P. simulans Zahlbr., nom. inval. non P. simulans Malme

P. pachythallina (Räsänen) Messuti syn: P. grisea var. pachythallina Räs.

P. papillulata Nyl. syn: P. pustulata var. erythrina Räs. syn: P. polycarpa Kremp. var. monospora Zahlbr. ?syn: P. idukkiensis Awasti & Srivastava

P. perrimosa Nyl. syn: P. subperrimosa Nyl. syn: P. basaltica Zahlbr.

P. pertracta Stirt. syn: P. gibberosa Müll.Arg. syn: P. woollsiana Müll.Arg. syn: P. virginea Müll.Arg. syn: P. moffatiana Müll.Arg. syn: P. nitidula Müll.Arg. syn: P. graphidioides Müll.Arg. syn: P. microspora Müll.Arg. syn: P. microsporella Zahlbr. nom. superfl.

P. pertusa (L) Tuck. syn: P. rupestris (DC.) Schaer. syn: P. pallidoflava Erichs. syn: P. leioterella Erichs. syn: P. colliculosa Körb. syn: P. communis DC.

P. pertusella Müll.Arg. syn: P. plicatula Müll.Arg. syn: P. straminea Müll.Arg. syn: P. communis var. pertusella Nyl. syn: "P. albidella Nyl." sensu Müll.Arg. syn: P. nigrescens Awasthi & Srivastava

P. petrophyes C. Knight syn: P. leucoxantha Müll.Arg.

P. phaeostoma Müll.Arg. syn: P. valdiviana Messuti & A.W.Archer

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. placentiformis (Mont. & Bosch) Müll.Arg. syn: P. communis var. placentiformis Mont. & Bosch syn: P. leioplaca var. placentiformis Müll.Arg.

P. pluripuncta Nyl. syn: P. gallica B. de Lesd. syn: P. gallica var. dealbata Llimona syn: P. rupicola (Fr.) Harm. var. bispora Werner

P. polysticta Vain. syn: P. ochrocarpa Vain.

P. porinella Nyl. syn: "P. albidella Nyl."[H-NYL 3826 pm, spores 95-112 µm; Sri Lanka] syn: P. ramuensis A.W. Archer & Elix

P. propinqua Müll.Arg. syn: P. torquata Müll.Arg. syn: P. marginata Nyl. ex Nyl. syn: P. rubescens Erichs.

P. pseudococcodes Müll.Arg. syn: P. subradians Müll.Arg.

P. pseudocorallina (Lilj.) Arnold syn: P. concreta Nyl. syn: P. ceuthocarpoides Zahlbr. syn: P. microsticta (Sm.) Erichs.

P. pustulata (Ach.) Duby syn: P. melaleuca (Sm.& Sowerby) Duby syn: P. canadensis Stirt. syn: P. limosa Zahlbr.

P. pycnophora Nyl. ?syn: P. scrobicularis Kremp.

P. pycnothelia Nyl. syn: P. leioplaca Schaer. var. dispora Müll.Arg.

P. pycnothelioides Vain. syn: P. "pycnothelia Nyl". sensu Müll.Arg.

P. quartans Nyl. syn: P. ikomae Yasuda ex Räs.

P. quasiae (Fée) Nyl. syn: P. leioplaca var. quassiae Nyl.

P. reagens (Zahlbr.) Redinger syn: P. reagens var. vegetior Redinger syn: P. tuberculifera Nyl. var. reagens Zahlbr.

P. rhexostoma Nyl. ex Hue syn: P. finkii Zahlbr. ex Fink

P. rhodiza Nyl. syn: P. gracilis Müll.Arg. syn: P. albidella Nyl. var tetramera Müll.Arg.

55 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. rigida Müll.Arg. syn: P. aberdarensis Dodge syn: P. rosacea var. octospora Zahlbr.

P. rochae Räsänen ?syn: gibbosa H. Magn. ?nom inval. syn: P. glebosa Räs. nom.inval. non P. glebosa Müll.Arg.

P saximontana Wetmore syn: P. christae Dibben & Poelt

P. simoneana A.W.Archer & Elix ?syn: Hämet-Ahti 3291a (H) Pertusaria sp. ex Argentina

P. sommerfeltii (Flörke) Fr. syn: P. melastoma Nyl., Flora 47: 489 (1864) [not in Zahlbruckner's Catalogue]

P. sorodes Stirt. syn: P. subcommunis Nyl.

P. subcorallina Nyl. ?syn: P. arsenei de Lesd.

P. submultipuncta Nyl. ?syn: P. sanguinulenta Zahlbr

P. subobducens Nyl. syn: P. tuckermanii Erichs.

P. subobductans Nyl. syn: P. glauconitens Müll.Arg. syn: P. platypora Müll.Arg. syn: P. fuscella Zahlbr. syn: P. kurohiensis Vain. syn: P. subobductans var. cinereoglauca Vain. syn: P. commixta Vain. syn: P. conferta Vain. syn: P. rugulosa Vain. syn: P. sublaeviganda var. pauperior Vain. syn: P. subrugulosa Zahlbr. syn: P. pachyplacoides Räs.

P. subochracea Stirton syn: P. hakkodensis Yasuda ex Räsänen

P. subrigida Müll.Arg. syn: P. depressa (Fée) Mont. & Bosch var. octomera Müll.Arg. syn: P. leioplaca var. octomera Müll.Arg.

P. subrosacea Zahlbr. syn: P. subrosacea var. evolutior Zahlbr.

P. subventosa var. ventosa Malme syn: P. paeminosa A.W.Archer

P. subverrucosa Nyl.; syn: P. callispora Zahlbr.

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Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. tazzekensis Werner ?syn: P. aceroae C.Hdez.-Padr, Etayo, I. Perez-Vargas & Elix

P. tejocotensis de Lesd. syn: P. arizonica Dibben

P. tesselaria Müll.Arg. syn: P. columnaris Malme syn: P. victoriana Lamb

P. tetrathalamia var. decipiens Müll.Arg. syn: P. leioplacoides var. decipiens Müll.Arg. syn: P. leioplacoides var. enucleans Müll.Arg. syn: P. tetrathalamia var. enucleans Müll.Arg.

P. texana Müll.Arg. syn: P. disticha Erichs. syn: P. obsolescens Nyl. syn: P. matsuzawae Yasuda ex Räs.

P. thiospoda C. Knight syn: P. leiotera Müll.Arg. syn: P. minuta C. Knight syn: P. subrugosa Nyl. syn: P. schizostomella Müll.Arg. syn: P. antillarum Vain. syn: P. stramineocarnea Vain. syn: P. intermedia Vain. syn: P. simplicata Vain. syn: P. xanthodes var. Vain. syn: P. xanthodes var. stramineoalbida Vain. syn: P. megapotamica H. Magn. syn: P. montevidensis Zahlbr. syn: P. straminella Malme

P. thwaitesii Müll.Arg. syn: P. wilsonii A.W.Archer

. P. trachythallina Erichs. syn: P. laevigata (Nyl.) Arn. var. discrepans Erichs.

P. trimera (Müll.Arg.) A.W.Archer syn: P. pustulata (Ach.) Duby var. trimera Müll.Arg. syn: P. melaleuca (Sm. & Sowerby) Duby var. trispora Müll.Arg.

P. tropica Vain. syn: P. pulchretincta Zahlbr. Indonesia; ?syn: Lepra hypothamnolica (Dibben) Lendemer & R.C. Harris [P. hypothamnolica Dibben]

P. truncata Kremp. syn: P. ascripta Stirt. syn: P. fumosa C. Knight syn: P. cupularis C. Knight, syn: P. subglobulifera Nyl. syn: P. nothofagi Zahlbr.

P. trypetheliformis Nyl syn: P. leioplaca var. trypetheliformis Nyl.

57 Archer & Elix, World-wide key to Petrusaria (including Lepra), Aug. 2018

P. tuberculifera Nyl. syn: P. tuberculifera statu isidiomorpha Nyl.

P. variolosa (Kremp.) Vain. syn: P. subvaginata f. variolosa Kremp. syn: P. subfallens Vain. syn: P. velatula Vain.

P. velata (Turner) Nyl. syn: P. chilena Zahlbr. syn: P. pulchella Malme; syn: P. skotsbergii Zahlbr. syn. P. commutans Vain. syn. P. subvaginata Nyl syn. P. albovelata Zahlbr. syn: P. chilena Zahlbr. syn: P. santamonicae Dibben syn: P. secedens Zahlbr. syn: P. velatoides A.L. Sm. p.p. syn: P. multipunctata var. colorata Zahlbr. ?syn: P. velata subsp. sublaevata Nyl. [thallus K-ve, C+red, KC+red;] syn: P. speciosa Høeg

P. velloziae (Vain.) Erichs. syn: P. velloziae (Vain.) A.W.Archer & Elix , nom. inval. syn: P. tuberculifera Nyl. var. velloziae Vain.

P. wattiana Müll.Arg. syn: P. wattiana var. fulvescens Müll.Arg.

P. xantholeucoides Müll.Arg. syn: P. moreliensis de Lesd. US syn: P. coccopoda Vain.; Philippines syn: P. kinigiensis A.W.Archer et al. Africa

P. xanthoplaca Müll.Arg. syn: P. persulphurata Müll.Arg. syn: P. sulphurata Müll.Arg. (1893) non P. sulphurata Müll.Arg. (1889) syn: P. citrina A.L. Sm.

58